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THE  NAUTILUS 


Volunw  116,  Niiiuhcr  1 
April  4.  2002 
ISSN  0028-1344 

A  quart erhj  devoted 
to  vuilacolo^ij. 


voocis  Ho:-' ; 


APR  1 


Wooes  t-ioc,  ij.M  02043 


EDITOR-IN-CHIEF 

Dr.  Jose  H.  Leal 

The  Baile\-Mattliews  Sliell  Museum 
3075  Sanibel-Captiva  Road 
Sanibel,  FL  33957 

MANAGING  EDITOR 

Christina  Petrikas 

The  Bailey-Matthews  Shell  Museum 
3075  Sanihel-Captixa  Road 
Sanihel.FL  33957 

EDITOR  EMERITUS 

Dr.  M.  G.  Harasew'Ych 

Department  of  Invertebrate  Zoolog)' 

National  Museum  of 

Natural  HistoiT 
Smithsonian  Institution 
Washington,  DC  20560 

CONSULTING  EDITORS 
Dr  Riidi'j;er  Bieler 
Depart ment  of  ln\ertebrates 
Field  Museum  of 
Natural  Histon- 
Chicago.  I Lfi0605 

Dr.  Arthur  E.  Bogan 

North  Carolina  State  Museum  of 

Natural  Sciences 
Raleigh,  NC  27626 

Dr.  Philippe  Bouchet 
Laboratoire  de  Biologic  des 

Invert^bres  Marins  et  Malacologie 
.Museum  National  dTIistoire  Naturelle 
55,  rue  Buffon 
Paris,  75005  France 

Dr  Robert  Il.Covvie 

C;enter  forCon.servation  Research 

and  Training 
Uni\ersilvof  llawaii 
3050  Maile  Way,  Ciilmore  409 
Honolulu,  HI  96S22 

Dr  Robert  T.Dillon.  Jr. 
Department  of  Biology 
College  of  Charleston 
Charleston.  SC  29424 

Dr.  Eileen  H.  Jokinen 
S234  E.  North  Shore  Road 
SaullSte.  Marie,  MI  49783 


Dr  Douglas  S.  Jones 
Florida  Museum  of  Natural  History' 
Universit)-  of  Florida 
Gainesville,  FL  32611-2035 

Dr.  Harry  G.  Lee 

ISOl  Barrs  Street,  Suite  500 

Jacksonville,  FL  32204 

Dr  Charles  Lydeard 
Biodiversit\-  and  S\stematics 
Department  of  Biological  Sciences 
University  of  Alabama 
Tuscaloosa,  AL  35487 

Dr.  James  H.  McLean 
Department  of  Malacology 
Los  Angeles  C'ounty  Museum  of 

Natural  History 
900  Exposition  Boulevard 
Los  Angeles,  CA  90007 

Dr  Paula  M.Mikkelsen 
Department  of  Living  Invertebrates 
The  American  Museum  of  Natural 

History 
NewYork,  NY  10024 

Dr.  Diarmaid  6  Foiglii! 

Museum  of  Zoolog)' and  Department 

ol  Biology 
Universitv  of  Michigan 
Ann  Arbor  MI  48109-1079 

Dr  Custa\  Paulay 

Florida  Museum  of  Natural  History 
Uni\ersit\-  of  Florida 
Gainesville,  FL  3261 1 -2035 

Mr  Richard  E.  Petit 

RO.  Box  30 

North  Myrtle  Beach,  SC  29582 

Dr  Gar)-  Rosenberg 
Department  of  Mollusks 
The  .Academy  of  Natural  Sciences 
1900  Benjamin  FranklJTi  Park-\vav 
Philadelphia,  PA  19103 

Dr  Angel  Wikles 
Department  of  .Malacology 
Los  Angeles  Count)  Museum  of 

Natural  Ilistoiy 
900  Ivvposilion  Boulevard 
Los  Angeles,  CA  90007 


Dr.  Geerat  J.  Vermeij 
Department  of  Ceolog)' 
University  of  California  at  Davis 
Davis,  CA  95616 

Dr  G.  Thomas  Waiters 

.Aquatic  Ecology  Laboratoiv 
1314  Kin  near  Road 
Columbus,  OH  43212-1194 

Dr.  John  B.  Wise 

Houston  Museum  of  Natural  Science 

Houston,  TX  770.30-1799 

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THE  €7  NAUTILUS 


CONTEXTS 


\'()Imiic  116,  Niunlx'i'  1 

April  4.  2002 

ISSN  002H-1344 


Eugene  \'.  Coan  RfCt-iit  castfrn  Pacific  species  ol  SiiiiLiiiiiuildrid  and 

Psrnuiilillti  i  BixaKia:  Psamoliiidae ) 1 

^^'allel•  Narclii  The  aiiatoiin  and  hinctional  ni()i"pli()I(>i;\  ol   Tu  rhi 

Fabio  di  Dario  vciitnci>\ii  (Ciraw  1S.3S)  (BixaKia:  X'cncridac) 13 

Berntl  R.  Sehone  Usi>  nl  ilNIDS  ilif\ainctli\ldisila/anc)  to  dn  oii;anic 

David  Bentlev  microstnictures  in  etclictl  hi\al\i'  iiiolliisk  and  lianiat'lc 

shells 25 

Philippe  Bouchet  Gone  with  the  wind:  a  pelaij;ic  marine  spi'cies  dcsci-ihed  as 

an  endemic  land  snail  from  the  Bahamas 32 


Note 

Miehael  Rex  Bioo;eoy;raph\  of  the  ileep-sea  i^astropotl  I'lihizz'ui  pliiiiorhis 

(Dall.  1927):  an  micominon  form  ol  rarit\ 36 


Notiee 24 


'I'liis  piililic-atidii  is  sponsored  in 

pai'l  l)\ 

tile  Stall'  ol  I'loi'ida,  Dcpartnu'iit 

of  State. 

Dixisiou  ol  (Cultural  Ailaiis, 

and 

tlu'  Florida  Arts  Coiiucil 

THE  NAUTILUS  llfii  11:1-12.  2002 


Page  1 


Recent  eastern  Pacific  species  of  SangiiuioJaria  and  Psammotclla 
(BivaKia:  Psammobiidae^ 


Eugene  V.  Coan' 

Department  ol  Invertebrate  Zoolotn.' 

and  Geolog)' 
Caliiomia  Academy  ol  Sciences 
San  Francisco.  CA  94118-4599  USA 
g;ene.coan@sierrachil).org 


ABSTRACT 

There  are  three  Recent  eastern  Pacific  species  ol  Sdii^iiiiinlcir- 
ia,  S.  ovalis  Reeve,  1S57.  .S,  tcUinoides  A.  Adiuns.  1S5(),  and  .S. 
tenuis  Olsson,  1961,  and  one  species  (if  P'idiniiKitclla.  I'  hciiini 
(Pilsbn  and  Lowe,  1932).  A  neot\]i('  lor  /'  hcrliiii  and  Iccto- 
hpes  lor  S.  tellinoklcs  and  its  junior  s\iion\ni  S'.  purpurea  are 
designated  herein.  The  distributions  of  the  species  are  docu- 
mented, along  with  their  fossil  occurrences  and  their  relation- 
ships to  other  Recent  and  to  fossil  species. 


INTRODUCTION 

Ha\ing  discussed  the  eastern  Pacific  repi'esi'utatixcs  ol 
tlie  genera  Hvtcrodonax  (Ct)an,  1973:  46—46)  and  Gnri 
(Coan,  2000).  I  herein  complete  the  rexiew  ol  tlie  lamilv 
Psammobiidae  with  treatment  ot  four  speties  tliat  have 
previously  been  assigned  to  tlie  genus  Saiij^uiuol/iriu. 

Previous  reviews  of  San^uinolaria  are  those  ol  Reeve 
(1857)  and  Beitin  (1880).  Trv'on  (1869)  listed  tlie  then- 
known  species.  Dall  (1898,  1900:  978-979)  and  Wilhui 
(1993)  di.scu.ssed  the  genera  of  the  Psammobiidae. 

Thus  tar.  there  are  no  papers  on  the  anatnmv  or  bi- 
ologv  of  S(i)i^ui)iolaria  or  Fsaiuinotella. 

M.ATLRIALS  AND  METHODS 

In  tlie  following  treatment,  eacli  valid  ta\on  is  followed 
bv  a  sviionvniv.  information  on  tApe  specimens  and  tvpe 
localities,  notes  on  distribution  and  habitat,  iind  an  ad- 
ditional discussion. 

The  svnonvmies  inclutle  all  major  accounts  about  the 
specie.s,  but  not  most  minor  mentions  in  the  literature. 
The  entrie.s  are  arranged  in  chronological  order  under 
each  species  name,  with  changes  in  generic  allocation 
from  the  previous  entn.  if  anv,  and  other  notes  given  in 
brackets. 

'  Mailing  aildress:  891  San  Jude  .Venue.  Falo  Alto.  CA  94.)()(v 
2640,  US.A;  also  Research  .\ssociate.  Santa  Bad)ara  Museuni 
of  Natural  Histon'  and  Los  .\ngeles  Countv  Musenni  ol  X.it- 
urai  HistoA. 


The  distributional  inlormation  is  based  on  Ht'cent 
specimens  I  have  examined,  except  as  noted.  Habitat 
information  is  scant,  because  most  material  has  been  col- 
lected in  beaclidrift.  Fossil  occiuTenees  are  taken  from 
the  literature. 

References  are  provided  in  the  Literature  Cited  lor 
all  works  and  ta\a  mentioned. 

Moi-jjhological  Characters:  -\  combination  of  shell 
shape  and  color,  .uid  various  aspects  of  the  shape  oi  the 
pallial  sinus  sidfice  to  distinguish  among  the  four  species 
discussed  here:  these  arc  detailed  in  the  tlescrijitions  and 
in  Table  1. 

Abbre\iali<)ns:  The  following  abbreviations  are  used 
in  the  text:  ANSP,  .Academv  of  Natund  Sciences  of  Phil- 
adelphia, Philadelphia,  Penn.svlvania,  USA;  BMNH, 
British  .Museum  (Natural  Iliston)  collection.  The  Nat- 
ural Iliston-  Museum.  London,  England;  C.\S,  (.'alilor- 
nia  Academv  of  Silences,  San  Francisco,  California, 
USA;  ICZN,  International  C!onnnission  on  Zoological 
Nomenclature;  L.\{'M.  Natural  Iliston  Museum  ol  Los 
Angeles  Countv.  (iaiifornia.  USA;  PRI.  Paleontological 
Research  Institution,  Ithaca.  New  York.  USA;  MCZ. 
Museuni  of  Comparative  Zoologv.  Hanard  Uiiiversitv. 
Cambridge,  Massachusetts.  USA;  MNHN,  Museum  na- 
tional d'llistoire  luiturelle.  Paris.  France:  SHMNH.  S;m- 
ta  Barbani  .Museum  of  Natural  ilistorv.  S.inla  Bariiara, 
Calilorni;i.  USA;  SDMNII,  S;in  DiegoMnseum  of  Nat- 
ural Ilisloi-v,  S;ui  Diego.  California.  USA;  UMML.  Uni- 
vcrsitv  of  Miiuni  \huine  Laboraton-.  Rosenstiel  School 
of  .Miirine  and  .\tmospheric  Science.  Miami,  Florida, 
USA;  USNM.  United  St;ites  National  Museum  collec- 
tion, NatioiKil  Museum  of  N;itural  llistorx',  Smith.sonian 
Institution,  Washiuglou.  DC.  US.\;  Z.MC;,  Zoologisk  Mu- 
seum Copenlnigen.  Denmark. 

.\hilciial  in  the  private  collections  ol  Carol  C.  Skog- 
huul.  Phoenix.  .\ri/ona,  US.\;  anil  Kirstie  L.  Kaiser, 
Puerto  \  ;illarta,  Jali.sco,  Mexico,  was  also  examined. 


Pagf 


THE  NAUTILUS.  Vol.  IIH.  \,..  1 


Table  1.    Kc\  (lilTrR'ntiatins;  cliaractcrs.  si/r.  ,iikI  tn'i|iiciic\  (il  Ea.stcni  Pacific  Sini'j_ini)i<liirin  and  Psiiuituofcllii 


Color 


Shape 


allial  smiis 


MiLKimum        No.  lots 
.size,  mm         studied 


S.  OKulls 

S.  tcUUioUlcs 
S.  tenuis 

P.  hciiini 


white,  with 
beaks 

purplish  rci 

while 

purplish  re( 


pink      etjuivalve,  ovate,  equilateral 


e(jiii\al\e.  oNate-elonsjate. 

Ioniser  posteriorK 
e((ui\al\'e.  o\ate-elouijate, 

longer  anIeriorK' 

right  \aKe  more  iiiHated. 
elongate,  lontrer  posteriorK 


gieatk  expanded,  pointed 

dorsallv,  meets  pallial  line  at 

appro.x,  50°  angle 
expanded  dorsallw  meets  pallial 

line  at  90°  angle 
greatK'  expanded,  pointed 

dorsallv,  meets  pallial  line  at 

approx,  75°  angle 
elevated,  rounded  dorsalK,  meets 

pallial  line  at  approx.  .30°  angle 


34 


20 


72 

79 

34 

1 

93 

98 

studieil: 

198 

.SYSTEMATICS 

FamlK'  Fs;inini()!)iidae  Fleming.  1S2S 
C.eiius  SaiiiiuinohirUi  Lamarck,  1799 

Sanguiiudaiia  Lamarck.  1799:  S4. 

Tvpp  .species  (li\    m()uot\p\  ^;      Soltii  sdii'j^iiiiiohnlirs  (imelin. 
1791:  3227,' 

Dt'scriplion:  lv|ui\aKi'.  with  a  narrow  posterior  gape. 
Periostraciiin  (liin.  Pallial  siiurs  deep,  mocleratelv  to  ven' 
elexatcd  dorsalK,  its  doisal  line  with  an  expanded  nniscle 
attachment  area  just  anterior  to  posterior  atlduetor.  Pos- 
terior enieiform  mnscle  scar  without  a  small  anterior  sat- 
ellite scar  Hinge  n.irrow  to  moderate  in  width:  teeth 
small;  n\  mnh  weak. 


Sau<iui)U)laria  oralis  l-leexf. 
l'"ignres  1.  2.  9 


hS57 


Saiiiiuiuiilnria  ill  alis  Hee\e.  i  S57:  pi.  1.  fig.  2;  Nhircli.  hSfid: 
185;  CariXMiter,  18«4:  563  [LS72  reprint:  49|  l;is  a  possihh> 
suionym  ofS'.  inininla]:  Tivon,  1869;  78;  Bertin,  h880:  84; 
iball.  1898;  61  |as  a  s\iionv'm  ol'S.  tcllinoide.'ih  Keen,  1958; 
188,  189,  fig.  460  |as  a  separate  species):  Keen.  1971;  243. 
(ig.  610,  244  [not  to  be  confused  with  lliuliihi  otalis  Ber- 
lin. 1880;  92,  pi.  4,  fig.  5a,  h,  a  species  ol  Solrlilliiui  de- 
.scrihed  from  -.m  unknown)  loealit\|. 

S<tnii,\iin<ilana  vcspciiiiui  Pilshiv  and  Lowe.  1932;  90-91.  I  II 
[;ls  "Si-iiiclc"  Vf.spciiina].  pi.  12.  figs.  3,  4;  llertkin  ;ind 
Strong,  1950;  220;  Keen,  1958;  188  |as  a  .sviiouym  of  ,S. 
(HY/Z/.s]:  Ols.son.  1961;  ,349,  558;  pi.  85.  fig.  5'fas  a'sep;n-ate 
.species];  Keen,  1971;  244  [as  a  svnon\ni  of  S.  (iiiilis\: 
Men/    1986:  41. 

Description:  Ovate.  etjuivaKc.  linn,  approxiuuiIeK" 
efinilateral  (heaks  at  4S-50'7c  I'rom  ;mlerior  end);  an- 
terior end  ronnded;  posterior  end  aeutelv  ronnded. 
without  a  radial  sniens.  Pallial  sinus  large,  much  ex- 
tended and  pointed  dorsallv,  meeting  pallial  line  at  an 
appro.ximately  5()-  augl..  prochicing  a  short  extent  of 
noii-fonHnence  l  Fig.  9)  Sculpture  of  fine  eommarginal 
striae.  Color  pink  iicr  uniliunes.  heeoniing  white  to- 
ward ventral  margin;  color  \isihle  botii  cxteriorK'  and 
interiorly.  Length  to  .34.2  mm  |PiU  25921;  Ciil'mico. 
Panamal. 


Type  Material  and  Localitie.s: 

S(in<s,iiinol<iriii  ornlis — BMNI 1  1957  7  15. 1 .  holotxpe.  with  bro- 
ken right  valve;  length,  22.2  nun;  height.  14.0  mm;  width 
(left  valve),  3.1  mm  (Fig.  1).  "CJeutral  .Vmerica",  Hugh 
('uniing  The  localitx  is  here  elarified  as  being  San 
Juan  del  Siir,  Rixas  Province,  Nicaragua  11.3°N!.  tol- 
lowing  KJZN  Cixle  Recommendation  76A.1.4. 

Sinifj,iiiiu>liiria  vc.spt'iiina — ANSP  155013.  holot\pe,  pair; 
length,  31.1  nnn;  height.  20.9  nnii;  width,  1L3  mm  (Fig, 
2).  ANSP  .398873,  paratype;  length,  26.4  mm.  SDNHM 
50773,  paratvpes,  6  pairs,  2  vahes.  San  Juan  del  Sur,  Rivas 
Province,  Nicaragua  (1L.3°N);  H.N.  Lowe,  193L  ANSP 
154663,  paratvpe,  left  \al\e;  length.  29.7  mm.  Corinto, 
Chinandega  Province,  Nicaragua  (12..5°N);  H.  N.  Lowe, 
1931. 

Distribution:  (;u;i\ui;is.  Som)ra.  .Mexico  (27.9°N) 
[CAS  1.54.3fi9],  to  (aianieo,  Los  Santos  Province,  Panama 
(7,2°N)  [PHI  2.5921];  from  the  intertidal  zone  to  .37..5  m. 
.Material  examined:  20  lots. 

Material  trom  the  Golfo  cie  Panama  that  has  been 
misidentified  as  this  species  [USNM  962S.3.  962S7. 
96353,  9fi.36L  96383],  or  labeled  as  Siin<:,iiini)liin(i  s/j. 
[Kaiser  collection],  while  eloselv  resianhling  a  Saiigiiin- 
olarid.  Ikis  conspicuous  lateral  teeth,  a  low,  elongate  pal- 
lial sinus,  and  slightlv  oblique  connnarginal  seuljitnre. 
This   material   is   instead   Tclliiia    [Utiiclliiia^   uinu/diid 


Ihrllcm  ,111(1  Strong.  1949  (pp.  S.5-S6.  97 
26  i  I  see  ;ii,so:  Ols.son.   1961;  409). 


il.   l.f 


1gS. 


Discussion:     This  sjiec  ies  is  vei"v  similar  to  its  western 

.Miaulic    I lologue.    .S.    stin<nii\ioicnt(i   (Cwiielin.    1791: 

322.5 — as  Solent  |svnon\ins;  Solcii  fiiiYilti.s  S[)engler. 
1794:  111:  Tclliiia  lihucuiula  Hciding.  179S:  1S6:  .SV;ii- 
iiuiiiolaiid  rosea  l.:uiiai'ck.  ISOl:  12.5:  Lt'liaria  rosacea 
Schumacher.  1S17:  122-12.3,  pi.  6;  Saniiniiiolana  iiirca 
.Morch.  IS5.3:  H)|.  which  occurs  from  Florida  and  Texas, 
through  the  WesI  Indies  lo  Hiazil.  Snti'^uiiiolaria  oralis 
diders  in  being  more  rounded  iuid  less  pointed  jiosle- 
rioiK.  in  h;i\ing  :i  n.trrower  hinge  pkile  and  liner  lunge 
teeth,  and  in  not  altaining  as  large  a  size  (S.  saiiiiiiiiio- 
Iciita  can  attain  at  least  4.'>.  I  mm  in  length  i.  .\ddition:illv. 
the  palli;il  simis  ol  .8.  saicj^iiiiiolciila  meets  liie  p:ilh.il  line 
;il  ;l  90"  ;ingle.  ;ind  il  does  not  rise  ,[s  hu"  dois:ilI\.  w  illinul 


E.  \'.  Coan.  200; 


Pacre  3 


Figures  1,  2.     S<nii^iiiiii>hiii(i  oidlis  Kccm 
Lowf.  Iciititli  31. 1  iiini. 


1.  ||iil(i|\pi'  (if  S    iir(;/(s,  l(-ii<;tli  22,2  nun.  2.  Ilcili)l\pr  ut  .S.  icspciiina  Pilshn  and 


a.s  shaqi  an  an<j;le  at  its  summit  (Fiii;iiri'  10'.  Tliis  wcstrrn 
Atlantic  species  niav  also  develop  tliickci-  slielK  lliau  an\ 
eastern  Pacific  material  of  .S.  oKilis  Si/irj^uhinliiriii  msca 
Lamarck  is  not  preoccupied  In  Sulcii  mscm  {imiliu. 
1791  (p.  3227).  which  was  based  on  a  (i'j;nie  in  ( .'heninit/ 
(1782:  pi.  7,  fig.  55)  that  seems  to  he  a  Si'lililliiia  This 
.species  was  attrihuted  to  the  Hed  Sea  l)\  Bertiii  1  ISSO; 
98).  It  is  also  not  preoccupied  In'  Tclliiui  ruM-ii  Cmelin. 
1791  (p.  3238),  which  is  based  on  an  illustration  in  Knorr 
(1771:  pi.  9,  fig.  3)  that  uia\-  be  of  a  specimen  ol  Asripliis 
dvjiomta  (Linnaeus.  1758:  rS87,  as  Wiuis). 

San^iiinohiriti  lifiva  Deshayes.  1855  (p.  32fii.  de- 
scribed from  an  miknowni  localit}'  (see  also  l^eexe.  1857: 
pi.  1,  fig.  1  I,  has  been  suggested  to  be  an  additiou:iI 
s\non\ni  of  .S.  sau^uinolcnta,  one  based  on  light-coloied 


material  iCosel.  19S9:  715h  (losel  based  this  conclusion 
on  two  lots  in  the  MXIIX  from  Ncnicrn/.  \cracru7,. 
Me\ic-o.  However  il'  material  in  the  Sli.MNII  (133229, 
345687)  and  the  (i.\S  (152575'  from  ni'ar  \eracruz  is 
correctK-  identified  as  S.  i  i/rcir  this  is  a  diiierent  species, 
indeed.  Diill  (  1898:  58)  gave  the  distribution  of  .S".  vilira 
as  being  from  Texas  to  ("olon.  I^anama,  but  (liis  needs 
to  be  verified,  'i'lic  Sl^\l\i!  and  (.'.\S  material  is  white 
autl  translneeut.  with  |);illi:il  sinuses  that  are  not  \ei"v 
dorsalK  extended  iiud  that  meet  the  pallial  line  almost 
veitieallv  I  LiL^ure  I  i  :  the  largest  specimen  is  52.4  nun 
in  leiiiith.  'Hie  Ivpi'  lot  iil  .S  1  Una  in  the  I^MNII  should 
be  examineil  to  be  certain  the  species  has  been  correctK' 
iiiteipreted. 

S(in<iiiiii(il<iiin    tiiirciuiiiclri    .Martens.    1S79    (p.    744) 


Pasie  4 


THE  NAUTILUS.  \ . 


Ifi,  \( 


[synonvni:  S.  africmia  Cosel.  19S9:  711-715;  pi.  1.  fig. 
K:  pi.  7,  figs.  26,  27)].  is  a  similar  West  .Mrican  species. 
(A  still  earlier  name  that  ma\  a]iply  to  tliis  species  is 
Tcllimi  arhatina  Spengler,  1798:  100.)  In  (iescrihing  ,S'. 
africana.  Cosel  (19S9)  differeiitiati'd  it  Irom  .S.  saufftin- 
olenta  as  l)eing  larger  and  more  elongate,  with  less 
brightly  colored  beaks.  These  characteis  would  also  sep- 
arate S.  mtrcot'incta  Irom  ,S'.  oiidis. 


San^uiiioliiria  tclUnoUlv.s  A.  .Ailanis,  1S50 
Figures  3-6,  12 

Siin<liiiri(il(iri(i  Iclliiu'iilrs  \.  .Xdaui.s,  1851):  170,  pi.  (i,  fia;.  (i; 
Kecxc,  1,S.57:  ])l.  1,  fig.  3:  CailH-nter,  IS57a:  286,  301: 
18571):  .31:  Morch,  1860:  185;  Caipeiitcr,  1864:  563  [1872 
reprint:  49J;  Tnon,  1869:  78  [as  "S.  lelliiiidcs"]-  Bertin, 
IcSSO:  84;  Dall,  1898:  58;  Meitlein  and  Strong,  1950:  219- 
220;  Keen,  1958:  ISS,  189,  fig.  462;  Olsson,  1961:  348, 
550,  pi.  77.  figs.  10.  11;  Keen,  1971:  244,  245,  fig.  611 

Tellina  miiiwlri  C.onU.  1851 :  90;  1853:  397,  pi.  16,  fig.  "l;  Gould 
and  Cai-peiiter.  1857:  199:  Caipenter.  1857a:'  226,  231, 
245,  301:  18571):  547  (in  Appendix  as  a  senior  ,s\iionvni  of 
.S.  piirj>iin'(i]:  Miircli,  1860:  185  [as  "muwacca"  and  a  svii- 
onvni  of  S.  icUinoidcfi];  Gould,  1862;  212;  Caqienter, 
1864:  537,  541,  543,  549,  563,  668  [1872  reprint:  23,  27, 
29,  35,  49,  154];  Tr\'on,  1869:  78  [as  a  svnonvni  of  ,S. 
iellinokh's];  Bertin,  1880:  84;  Dall,  1898:  16 '[as  a  .synonym 
of  .S.  tcUUwidcs]-  lleitlein  and  Strong,  1950:  219  [as  a 
s\iu)n\in  of  S.  Iclliiioides]-  Johnson,  1964:  110;  Keen, 
1971:  244  [as  a  s\non\ni  ol  .S.  Icllinoidcs] 

Sanguinolaria  puqwmi  Desha) es,  1855:  346;  Reeve,  1857:  pi. 
1,  fig.  5;  Gould  and  (Carpenter,  1857:  199  [as  a  svnonym 
of  S.  miniatii]:  Caipenter,  1857a:  226,  231,  245,  301,  352; 
lS57c:  31,  548;  1864:  563  11872  reprint:  49];  Tiyon,  1869: 
78  [as  a  synonym  of  ,S.  frilinoidcs];  Dall,  1898:  61  [as  a 
synonym  of  .S.  trlliiioidcs]:  llertlein  and  Strong,  1950:  219, 
251,  pi.  2,  figs.  5,  8  [as  a  separate  species];  Keen,  1958: 
188,  189,  fig.  461;  Keen,  1971:  244  [as  a  .synonym  of  .S. 
Icllinoidc's] 

Sanffiiiu>hiriu  lellinoidc.s  cloufidtd  Morch,  1860:  185;  llertlein 
and  Strong,  1950:  220  ]not  preoccupied  1)\-  Ciiri  (Pstiiii- 
moUicna)  clonfialn  (Lamarck,  1818:  514 — as  Vs(nnmohia), 
which  is  widespread  in  the  Indo-Tacific  (Willan,  1993:  61- 
64)] 

Description:  Ovate-elongate,  equivalve,  some\\h;it 
tliicker-shi'lled  than  ,S.  ovalis  at  a  similar  si/.e,  becoming 
sturdy  in  large  specimens;  posterior  end  somewhat  lon- 
ger (beaks  at  40-42"t  from  anterior  end);  anterior  end 
roimded;  posterior  end  somewhat  produced,  set  oil  l)\ 
an  indistinct  radial  sulcus  a[)proximatelv  Kvo-thirds  dis- 
tance Irom  end  generally  most  evident  in  large  speci- 
mens. Pallial  simis  large,  proiluced  and  pointed  dorsallv, 
meeting  pallia!  line  at  90'  anule,  its  ventral  margin  thus 
completelv  confluent  with  pallial  line  (Figure  12).  Sculp- 
ture ol  fine  ccuiniargiiial  striae,  (.'nicilbrm  muscle  sears 
inflated  in  large  spcciin.Mis  I.  ;ferior  color  pink  to  pur- 
ple, sometimes  with  d;nkcr  curnmarginal  color  bauds: 
interior  often  dark  ]-iuiplish-re(l  Length  to  71.5  mm 
(ANSP  220.326:  Aeapiilco.  C.uerrero.  Mexico). 


Type  Material  and  Localities: 

Siiitiiuiiiolarin  liUinoidcs — B.VLMl  1966540/1,  pair  lectotype 
here  designated;  length.  32.6  nun;  height.  19.2  nun: 
widdi,  8.4  mm  (Figm-e  3).  BMNH 1966540/2,  paralecto- 
t\pe;  length,  32.5  mm.  Gulf  of  California.  The  locality'  is 
here  clarified  as  being  GuaMiias,  Sonora,  Mexico 
(27.9°N).  loHowing  ICZN  Code  Recommendation  76A.1.4. 

Tellina  minUitd — .\1C;Z  169258,  holotype,  pair;  length,  51.0 
nun;  height,  33.0  mm;  width,  14,3  mm  (Figure  4).  San 
Juan  [del  Sur,  Ri\as  Province,  Nicaragua]  (11.3°N);  Lieut. 
T.  P.  Green.  The  localitv  was  mistakenK  given  hv  Johnson 
(1964:  110)  as  being  in  Orange  County,  Califonii:i, 

Stnii^iiiiKildiiii  imrpiircd — B.VINII  19f)6539/l,  leclotvpe  here 
designated,  pair;  length,  50.0  nnn;  height,  31.7  nun; 
width,  15.1  mm  (Figure  5).  BMNH  1966539/2,  3,  para- 
lectotypes,  pairs,  lengths,  48.4  mm,  44.4  mm.  The  original 
specimens  came  from  the  collection  of  Hugh  Cuming,  but 
the  t\pe  loealit\  was  gi\en  as  unknown  and  is  here  clar- 
ified as  being  Guaymas,  Sonora,  Me.vico  l27.9°N),  fol- 
lowing ICZX  Code  Recommendation  7(i\.1.4. 

SdUf^uiiioldiia  tellinoides  elon^dld — ZMC  unnumbered,  holo- 
tvpe,  pair;  length,  52.4  nnu;  height,  30.2  nnn;  width,  13.3 
nnn  (Figure  6).  "Realejo"  [Corinto,  Chinendega  Province, 
Nicaragua]  (12.5°N);  A.  S.  Oersted,  1846-1848. 

Distribution:  Punta  Pescadero,  Pacific  coast  of  Baja 
Calilornia  Sur  (23.3°N)  [USNM  22964],  into  the  Golfo 
de  California  as  far  north  as  Punta  Diggs,  Baja  Cidifornia 
(30.9°N)  [CAS  150381],  and  Puerto 'Libertad,  Sonora 
(29.9°N)  [ANSP  184183],  Mexico,  to  Cojinu'es,  Esmer- 
aldas  Province,  Ecuador  (0.4°N)  [PRI  259208];  intertid- 
a!  /one  to  14  m.  Material  examined:  79  lots.  San^uitio- 
laria  tellinoides  Ikus  been  reported  from  the  Pliocene 
|ania  Formation  at  Puerto  Jama,  Manabi  Province,  Ec- 
uador (0.2°S)  (PiLsbn-  and  Olsson,  1941:  72).  and  (as 
"(iff.")  from  the  late  Miocene  Gatun  Formation  on  the 
Atlantic  coast  of  Panama  west  of  Colon  (W'oodring,  1982: 
673,  pi,  115,  fig.  12). 

Discussion:  The  pallial  sinuses  ol  this  species  become 
more  tlorsalK'  pointed  in  large  specimens.  This  species 
can  be  distiuguislied  from  i'saiitiiiotclla  heifini  in  being 
e(]ni\aKe.  and  1)\  its  more  rounded  outline,  less  atten- 
tuate  jiosterior  enil,  :ui(l  more  (l()is:il!\-  extentlcd  and 
pointed  pallial  sinus. 

Sdiiiiniiuiliiri/i  leiiui>:  Olsson.   1961 
Figni'c  7.  13 

Sdiipiiiioldiid  leiiuis  OLs.son,  IS)61:  .349,  558,  pi.  85,  fig.  6; 
Ke<'n.  1071:  244  [as  a  sviionvm  of  S.  (wnlis;  not  a  hom- 
ouMii  With  Psdiioiu'hid  tennis  Deshaves,  1855:  320,  a  ,svii- 
nuMii  111  ihc  I  IK  Id- Pacific  (.Idri  dnonudd  I  Deshaves,  1855: 
320— as  I'sdnwiohid)  (Willan,  1993:  22),  nor  with  Soletel- 
lirid  tennis  (Desha\es,  18.55:  .349 — as  Capsd  (Cdpsella)) 
from  the  I'liilippinr  Ishuuls  (Willui.   1093:  77)[ 

Description:  Ovate-elongati';  e({ni\aKe;  ant(>rior  (>nd 
somewhat  longer  (beaks  at  44'X  from  anterior  end);  an- 
terior end  rounded:  postciior  end  I'ounded.  with  :i  slight 
(race  ol  ;i  radial  sulcus  at  two-thirds  ol  wa\  to  postciior 
end.  Piillial  sinus  large,  exlendi'd  dorsalK',  meeting  ]iallial 
at   an  ■ij)pi'i)\iui:i(e  75"  angle  (I'^igure    13).   Scuiptnic  ol 


E.  \'.  Coan,  2002 


Pai^c  5 


Figures  3,  4.     SiniiS^iiinnldriti  ullininilcs  A. 
{ )i>ul(l:  li'iiiidi.  51 ,0  mill 


Adams.  3.  I  ,cilnl\pc  ol'  .S.  tclliiiniilcs.  lcii<j;tli  -i^fi  mm.  4.  Ilolotypc  (il  Tclliiia  iiiiiiialii 


fine  foiiniiai^iiial  .striae.  I'Alciini'  cnloi'  wliitc,  willi  a 
lit^lit  pink  railial  liaml  a  little  anterior  ot  midline;  while 
inteiiorK.  Lensjtli  t(i  oo.S  nnii  (li()l()t\pe). 

T>pe  Nraterial  and  Localily:  AXSF  21Si)il  lioh, 
t\pe.  pair:  leiiiitli.  3.1S  nnn:  lieit^ht,  26.1  inni:  widtii, 
10. U  nnn  ( Fii^nre  7l:  ( lanoa.  Manabi  Proxinee.  Kenadnr 
(0.5°N);  A.  A.  Olsson,  1958.  An  additional  fraiinientai-\ 
specimen  cited  h\  Olsson  (1961)  from  Pnnta  Montafiita. 
Gnavas  Province,  Ecuador  (1.8°.S).  would  he  a  paratypc; 


il  lias  not  been  located  in  tlie  r.MMI;.  I'nlortiinateK; 
none  ol  Olsson  s  specimens  ol  Suiiffiinolmid  lia\i'  \ct 
been  located  in  tlie  UMML  (\.  \'()ss,  e-mail.  2-3  May 
2001 1, 

Distnbiition:  Tims  far  known  onK  from  the  original 
specnneiis — ( .'anoa.  Manabf  Pro\ince  (0.3°\  *  ilioiotxpe), 
presnmabK  lo  i'lnita  .Montafiita.  (Jnaxas  Proxinee  ( 1.8°.S) 
(specimen  missing),  Ecuador.  Botli  specimens  were  col- 
lected in  beaclidrilt. 


Pa<'e  6 


THE  NAUTILUS,  Vol.  116.  NO    1 


Figure'.  .5,  (i.     Sdii-j^'iinnliinii  tclliiioidcs  A.  Adams.  5.  Lfctolvpc  (iT  .S',  imrpiircii   l)cslia\cs.  Icii'j;lh  oDd  niiii.  (>.  Il(il(il\]ic  nl  S. 
icllinoUlis  rlnu^aUi  NKircli.  linolli  .52.4  iiini. 


E.  \".  (Joan.  2002 


Figures  7,  8.     7.  Siiiii^iiiiHilariii  Icniiis  (iIssdii:  Imlnlxpc    liiiulli   >-vS  iiin 
Tcllina  hanlci/i  Bertiii;  leiit^tli.  fiT.o  mm. 


S.  I'sinitiiKitclld  IkiIiui     I'ilshiA   aiu!  I.iiwc  :  iu(il\  |ii' iil 


Page  S 


THE  NAUTILUS,  Vol.  116.  NO.  1 


Discussion:  Tliis  is  tlic  rare  Soiitli  Aiiit'rifan  alK  ot  S. 
oifilis.  It  cliflers  in  not  lia\inti  rose-tolored  l)eaks,  in  be- 
ing thicker  slielled,  and  in  having  a  more  rounded  pos- 


terior end. 


Genus  Psaminotclhi 


Icrniiannsen. 


1S.52 


Psammotella  Ik'rrniainiscn,  1S.52:  114,  c.v  "Fsaiiiiiiott'lle" 
Blainville,  1S28:  .541  | vernacular], 

T\pe  species  (bv  nion()t\pv):  'T.  nifcscens  Chenin.",  =  TcUina 
nifescens  Dillwvii,  ISIT:  85,  t'.v  Chemnitz  ins,  =  TclUna 
cntcnta  [Lightfoot],  1768:  10  [as  Tellina  "cmcntac"].  .58 
[as  T.  cntenta]'^:  first  reviser:  Rehder,  1967:  7.  Recent, 
western  .Vtlaiitic.  [mm  Psammotella  H.  .4danis  and  \.  Ad- 
ams, 1S.56:  39.3,  c.v  Deshayes  ms,  =  Psammotdlina  R-H. 
Fisciier,  1887:  1105,  new  name,  a  subgenns  o{  Can  and 
perhaps  a  senior  SNiion\ni  of  Psammotaena  Dall,  1900: 
976  (VVillan,  1993:  60)]. ' 

Description:  Shell  inequivalve;  right  valve  more  in- 
Hated;  posterior  gape  narrow.  Pallial  sinus  deep,  mod- 
erately elevated  dorsalK,  without  an  expanded  muscle 
attachment  anterior  to  posterior  atlductor  muscle  scar. 
Posterior  crncitorm  muscle  scar  with  a  small  anterior 
satellite  scar  Hinge  moderatcK  lieavA';  teeth  small: 
iniupli  weak. 

Discussion:  I  here  rank  this  ,Ni'w  World  taxou  as  a 
genus  because  of  its  major  differences  from  Sau^uhm- 
luria — conspicuous  differences  between  left  and  right 
\al\(>s.  plus  dillerences  in  its  pallial  sinus  and  cruciform 
muscle  scars. 

Psammotella  hcrtiiii  ^Pilsbn  and  Lowe,  1932) 
Figures  8,  14 

Tellina  hanlet/i   Bertin.    1S7S:   268-269   Inoii   Tellina  haiilei/i 

Danker  18.53:  53-54.  pi.  10,  figs.  4-6|. 
Sanfifiinolaria  heriini  Rilsbrv'  and  Lowe,  19.32,  new  n.inu'  lor 

Tellina  haulei/i  Bertin.  1880,  non  Dnnker   1853;  I'ilsbiv 

and  Lowe.  1932:  91.  )43.  pl.  10.  figs.  7.  8:  Hertlein  and 

.Strong,   19.50:  220-221;  Keen,   1958:   188,   189.  fig.  4.59; 

Olsson,   1961:  .349,  .5.50,  pi.  77,  fig.  8:  Keen,   1971:  244. 

245,  fig.  (5l2:  Hertz,  1986:  36. 
Tellina  nijesceiis  "Chenniitz",  auetl..  non  T  mfeseen-s  Dillwvn, 

1817,  ex  Chemnitz  ms;  Hanlev,  1846:  .307-308,  332.  pl. 

63.  fig.  213:  Caipenter,  18.57b:  32;  Rtimcr  1S72:  111-112, 

pl.  27,  figs.  1-3  [mm  Tellina  nifeseens  DilKwn.  IS17:  S,5, 

ex  (;licinnil/  iiisj. 

Description:  h^loiigale;  right  valve  conspicuously 
more  inflated  th.m  left;  posterior  end  longer  (beaks  at 
40-48'^  from  and  lit;;  .•■idi;  :uiterior  end  rounded;  pos- 
terior enfl  attenuated.,  set  off  bv  a  radial  sulcus  near  end 


in  riglit  \aKe,  tip  truncate;  posterior  end  siiuious  in  right 
vahe.  tip  suhtmncate.  Pallial  sinus  deep,  its  dorsal  mar- 
gin in  right  vaKe  ele\ated,  rounded  to  bhmtK'  angular; 
ventral  margin  of  sinus  meeting  pallial  line  at  an  ap- 
proximate 30°  angle,  confluent  with  pallial  line  for  ap- 
proximately 75%  of  its  length  (Figure  14).  Sculpture  of 
fine,  irregular  conunarginal  striae,  strongest  on  posterior 
slope;  right  \al\-e  with  radiid  striae,  strongest  along  \en- 
tral  margin  and  in  large  specimens.  Cniciform  muscle 
scars  bulbous  in  large  specimens.  Color  pink  to  puqile. 
sometimes  with  darker  conunarginal  bands.  Length  to 
92.8  mm  (LACM  71-179.12.  Punta  P.-qucfui,  Baja  Cd]- 
ifornia  Sur.  Mexico). 

Type  Material  and  Locality:  Tellina  lianlciji  Bertin 
was  based  on  the  figure  ot  Tellina  nifescens  "(Jhemnitz" 
in  Hanlev  (1846;  see  abo\e).  which  presumabK  came 
from  the  onl\-  localit\-  mentioned — Tunibes.  Tumbes 
Province,  Pei"u  (3.5°S).  This  specimen  has  not  been  lo- 
cated in  the  BMNH  (J.  Pickering,  e-mail,  7  June  2001). 
nor  in  the  Leeds  Museum,  present  location  of  a  sub- 
stantial ;unount  IIaule\s  material  (A.  Norris.  e-m;iil,  13 
)uue  2001).  The  specimen  selected  by  Pilsbrv  and  Lowe 
(1932)  as  iiolotvpe"  of  their  new  name— ANSP  152068 
from  ./\capulco.  Guerrero,  Me.xico  (16.9°N)  (Figure  S) — 
would  normalK  have  no  type  status,  because  the  txpe  of 
;i  renamed  junior  homon\ui  remains  the  original  tvpe  of 
that  taxon  (ICZN  Code  Art.  72.7).  Howex'er.' because  ( 1) 
Tellina  nifescens  was  long  used  to  refer  both  to  the  west- 
ern Atlantic  species  now  knowii  as  Psammotella  cnienta 
and  to  the  eastern  Pacific  /'  heiiini.  (2)  we  cannot  as- 
cert;iin  wliich  ol  these  two  species  Hanlev  actually  illus- 
trated, (3)  there  is  no  good  material  t)f  the  eastern  Pa- 
cific species  in  coll(>ctions  from  Tumbes,  Peni.  and  (4) 
PilsbiA  ;uul  Lowe's  "txpe"  is  ;i  fine  spetiinen  long  ;ic- 
ctjriled  t\pe  status,  this  specimen  is  here  designated 
neot\pe  of  Tellina  liaiilei/i  Bertin.  It  is  ;i  p;iir  of  \aKes 
measuring  67. .5  nun  in  length.  28. 1  mm  in  height.  ;uid 
12.4  nun  in  width. 

Distrihution:  L;ignn,i  ())n  de  Liebic  [Scinunons  La- 
goon], H;ij;i  California  Sur(27.8°\)  ]ANSP  22.5928],  La 
Paz,  Baj:i  C;iliforuia  Sur  !24.2°\)  [SBMNH  24586. 
24587],  ;ind  Kmpalnie,  Sonora  (27.9°N)  ]SBMNH 
1.35133],  Mexico,  to  M;ineor;i.  Tumbes  Province.  Peru 
(4.rS)  |CAS  1.54.370;  SBMXII  125767],  and  evidently 
as  far  south  ;is  (^)l;in.  I'iin;i  Province,  Peru  (5.0°S)  (Pa- 
redez  and  ( !;uxlo/o  ms;  Mnsco  de  llistoria  Natural, 
UTiivcrsickid  N;icioual  .\I;i\()r  de  S;in  .\I:ircos.  Lima. 
Peru);  inlerlickil  /one  to  17  ni.  \l,ileri,il  e\:nnined:  98 
lots. 


Figures  9-15.     Li:ii.',t      ,:■ 
scars  ol  left  and  righl  ■■.•'.         'I. 
(Gniehn):CAS  152576:  \h,i.   i, 
Veracmz,  Mexico;  lenL'th,  46  "  ; 
.S.  tennis  Olsson;  liolotvp":  liii'.'i!; 
length.  66.6  mni.  15.  P  <  nienta 


■■■■•a]  views  ol  slu'lls  ol  San^liitnohiria  ,un\  I'stnnninh  llii  sliewnig  liiii'j,!'.  palli;ll  sinns,  ;incl  .iildnetor 

■  -./'v  Reeve:  CAS  1.50375:  .\e,ipiileci   (aieirero.  Mexico:  length,  26,2  mm.  10.  S.  saii'diiinolenta 

'..u.iiha.  Bnr/.il;  length.  46,  I  hum    IL  ,S   litrea  Desliaves;  C.'^S  152576;  Boca  ik-1  j-lu).  \'ei-;icniz. 

\1.  .s   lellimmles  A.  Adiims;  .WSR  220.'i26;  .\c;i]inIeo,  Cnerrero.  .Mexico:  length,  41.5  nun,  13. 

■J  S  miM    M.  Psammotella  heiiini  iPilsbn  ;md  Lowe);  C.\S  1.50.380;  ,\capnlco.  Guerrero.  Mexico; 

.i'4lilloMf     .  (  olon.  P;in;iin;i  I'rovinee,  I'linam.i;  length.  (il.O  nnn. 


E.  \\  Coan.  2002 


Page  9 


10 


11 


12 


13 


14 


15 


Paire  10 


THE  NAUTILUS.  Vol.  116.  No.  1 


Discus.sion:  Psaiiiini>tvlla  licrliiii  is  \v\\  similar  (o  its 
western  Atlantic  lioniolos^ue.  /'  cnicnia  lll.iiilitloot]. 
17S6:  see  nncler  genus)  [.synonyms:  Tcllina  opcrciddta 
Gnielin,  1791:  3235  (as  "T.  opcrculalds"  in  Lugcluni 
printing);  Tcllina  nifcsceiis  nill\\-\n.  I  SI  7:  S5.  c.v  Chem- 
nitz ms].  The  last  sviions  m  is  naii  7'  rulrsrcns  (imelin, 
1791:  3238.  which  was  based  on  (;ualtieri  (1742:  pi.  25. 
fig.  C),  .seemingly  a  V<'ncnij)is.  perliaps  best  regarded  as 
one  of  the  nianv  s\non\iris  ol  V.  dectissatiis  (Linnaeus, 
175S:  690 — as  Venus),  as  suggested  bv  Carpenter 
(1857b:  32).  PsaininofcUa  rnicnta  occurs  throughout  the 
Caribbean  to  Hnizil.  PsuiniiioivlUi  rnicitta  dilters  from  P. 
bciiini  in  being  more  iuecjiiilateral.  with  a  llattener  left 
valve  and  a  more  inflated  right  \al\e,  and  a  less  dorsallv 
expandi'd  pallial  sinuses  in  both  \aKes  (Figure  15).  Oth- 
er supposed  dilTereiices  that  have  been  suggested  \an 
too  much  among  populations  to  be  useful.  For  example, 
Hertlein  and  Strong  (1950)  throught  tliat  /'  rnicniii  is 
narrower  posteriorlv,  but  this  does  not  seem  to  be  (he 
case  ifCnougli  lots  are  studied.  Thev  also  said  that  the 
pallial  siTiuses  ol  /'  cniciita  were  "more  arched  posteri- 
orlv" (?e.\tending  further  posteriorly)  and  "confluent 
with  the  pallial  line  lor  a  greater  distance."  but  I  laii  t 
see  nuich  difference  in  these  parameters. 

Psaininotclla  smitlncoodunnli  (Maur\-,  1917:  393-394 
[  =  229-230],  pi.  64  |  =  38|,  figs.  1,  2— as  San^iiinolaria 
{Psaiiiuu)t('lla)).  from  the  late  Miocene  Cercado  For- 
mation ol  the  Dominican  Hepublie  is  presuniabK  ances- 
tral to  these  two  species.  .Vlann'  pointed  out  its  affinits' 
to  P.  herliiti.  but  tlid  not  compare  them.  The  original 
figures  are  insulficientlv  clear  to  see  anv  differences 
from  either  Recent  species.  Weisbord  (1964:  372)  noteil 
that  this  Miocene  species  differs  from  Recent  material 
in  lacking  radial  striae  in  the  right  \aK('. 

Psammolclla  alottafii  Olsson  (1922:  432-433  |  =  260- 
261],  pi.  32  [  =  29].  figs.  5,  6 — as  S(iu<iiiiiioltiria  {Sam- 
motcUa)  |sic]  from  the  late  Miocene  Catun  Formation 
at  Banana  Hill  on  the  Atlantic  coast  of  Costa  Rica,  was 
biiseil  on  two  poorly  presenetl  \al\i's.  The  sjiecies  was 
said  to  be  more  elongate  posteriorK  than  /'  cniciilii  and 
to  lack  radial  striae  in  the  right  xaKc.  Weisbord  if964: 
372-373)  added  that  the  anterodorsal  margin  of  the  rigli( 
valve  and  the  posterodorsal  margin  of  the  left  \al\i'  were 
luore  concave  tiian  in  /'  cniciila.  Both  of  these  fossil 
species  are  tlins  far  (oo  |)oorl\  known  lo  draw  an\  useful 
conclusions. 

I'or  a  comparison  with  Saii'^iiinoldrid  Iclliiinidcs,  see 
undt-r  this  species. 

L.XCLUDKD  TA.XA 

Saii'^nhudari:!  (alifcrtiiaitd  Conrail.  1837  (p.  231.  pi.  17. 

fiii.  /  )  I.-,     ■liliirnirii.  atirlt..  lunn.  tiidi).  is  a  s\non\ni  of 

Mannr        r         ■!    (Liimaeus,     175S:    677 — as    Tclliiid) 

i^       •        •-■  •17). 

S-  (!■  p.  81),  is  now  regarded  as  a  s\ii- 

onvm  ol    •  ■':;  (C:oiuad,  1837:  230-231;  |il. 

17,  fig.  ()  :.    -^'K    I   426). 

S.  fli.sca  {Ha\.  :'''■     as  J  ,-,c/;/i»io/;/c/).  a  combiniilion 


ol  some  ;iutliors,  is  aSMloinni  of  Mucoiiui  hullliicd  (Lin- 
n;ieus,  1758)  (Coan  et'al..  2000;  417). 
.S.  <^rdiidis  Caipenter,  1857,  c.v  Gould  ms,  a  ikhiuh  uii- 
dtini  in  Caipenter  (1857a:  228,  349),  is  regarded  as  a 
prolnible  sviionvm  of  Xuttnllid  iiuttaUii  Conrad,  1837 
(Coan  et  aL,  2000:  426). 

S.  mitldlla  Conrad.  1837  (pp.  230-231;  pi.  17,  fig.  6)  is 
now  placed  as  Xnttdllia  iiuttidlii  (Conrad,  1837)  (Coan 
et  al..  2000:  426). 

S.  paciftca  (C^onrad.  1837:  241.  pi.  18.  fig.  13 — as  Psam- 
iitohid)  is  a  combination  bv  some  authors  for  the  species 
now  known  as  llvtvrodoiuix  pacificus  (('onr;id.  1837) 
(Coan  et  al.  2()()0;  428). 

S.  ndirnradidfd  Caipenter,  1860  (p.  1).  c.v  Xuttall  or 
(Conrad  ms.  is  a  iioiiicu  nudum  now  regarded  ;is  a  prob- 
able s\non\in  of  Gaii  cdlifornicd  (Clonrad,  1849;  121) 
(Coan'et  ;i[..  2000  426:  Coan,  2000:  3). 

ACKNOWI.EDCMENTS 

I  ;ippit'ciate  tlie  help  of  the  following  curators,  other 
personnel  and  their  institutions,  who  niaile  specimens, 
literature,  and  information  a\ailabk';  .^dam  J.  Baldinger, 
Museum  of  Comparative  Zoologw  Hanard  Uni\ersitv, 
Caiubridge,  Massaclnisetts,  US.\:  Ruck)  xon  Cosel,  Mu- 
sc'-um  national  d'Histoire  naturelle,  Paris,  France;  Lind- 
se\  T.  Groves  and  James  H.  McLean,  Natural  Histon 
.Vluseuni  of  Los  Angeles  Counts,  Los  Angeles,  ("alifor- 
nia,  USA;  Elizabeth  Kools,  Department  of  Inxertebrate 
Zoology.  California  Academv  of  Sciences,  Golden  Gate 
I\nk,  San  Francisco,  California,  US.\;  Da\id  R.  Lind- 
berg.  Museum  of  Paleontolog\,  Uni\ersitv  of  California, 
Berkelev.  ("alifonii;i,  USA;  Adrian  Norris.  Leeds  Mu.se- 
uiu,  Leeds.  I'aiglaud.  UK;  Joan  Pickering.  The  Natural 
Histoi-v  Museum,  London,  England,  UK;  T\juana  Nick- 
ens,  Natiouiil  Museum  of  Natural  Histon,  Washington, 
DC,  US,\;  GaiT  Rosenberg,  The  Acadeiu\  of  Natural 
Sciences,  Philadelphia,  Penns\i\auia,  US.A;  Paul  \'aleu- 
tich  Scott.  Sant;i  l^;irb;u';i  Museum  of  Natural  Histon. 
Santa  B;uiiara.  (!alilornia,  USA;  Ole  S.  Tendal  and  .\nnie 
Lone  \'e(lelsl)\.  /oologisk  Museum,  Copenhagen,  Den- 
uKirk;  \;me\  \'oss,  Uni\ersit\' of  .\liaiui.  .Miami.  Florida, 
ISA  ( !arol  ('..  Skiiglimd  and  kirstie  L.  Kaiser  gener- 
ousK  m;ide  ;i\;iil;ible  materials  or  information  from  their 
collections.  .\!;m  R.  K;ib;it  provided  some  ;ul\iee.  Lind- 
se\  T  (iroN'es.  Ke\in  L;nnprel!.  C;n()l  (,'.  Skoghmd.  Paul 
\alentieh  Scott,  ;uid  Richard  \\  illan  ukkIc  helpful  com- 
ments (III  llic  iiMiiiiscript.  SIku'oii  \\illi;mis  helped  pre- 
p;il"e  the  phUes. 

i.iii'Mail  BF,  CITFD 

Adams.  .\.  1850.  Dcsc  rijiliiins  of  new  species  ol  sluils  Iroiii  (lie 
("iiiiiiii>;iaii  cdllection.  I'roceeciings  ol  the  Zoologiciil  So- 
ciel\  (if  I. (.11(1(111  lor  1849  (200):  169-170,  pi.  6  |ripriiitecl 
in:  XiiiiaK  and  Ma'.;aziiii'  of  Natural  Histon  i2>7(37):  69, 
1S51I, 

Adams,  II  and  A.  .Aihinis,  1S56  [1S53~1S.")S|,  The  ueiicra  of 
l^ccciit  Miilliisca;  arraiiilcd  according  (o  tli(ir  iir(.iaiii/.a- 
tidii    \aii  \ii()rsl,  I.oihIiiii  liii).  3M— tl2,  Ncn.  1S.')6|. 


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ly the  property  of  the  Duchess  Dowager  of  Portland,  de- 
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Co Loudon,  viii  +  .3-194  pp.  [concerning:  Rehder, 

1967] 

Linnaeus,  C.  1758.  S\s(ema  naturae  per  regna  Ilia  natiu'ae  .  .  . 
editio  decima,  rcformata.  \'ol.  1.  ,\nimalia,  L,  SaKii.  Stock- 
holm, 824  +  iii  pp. 

Martens,  K,  E.  von.  1879.  Ubersicht  der  son  Herru  W.  Peters 
von  1843  bis  1847  in  Mossambi(jue  gesannnelteu  Mollus- 
ken.  Monatsberichte  der  Koniglicheu  Preussisclieu  .\ka- 
demie  der  Wissenscliaften  zu  Berlin  for  1879:  727-749. 

Maury,  C.  J.  1917,  Santo  Domingo  t\pc  sections  and  lossils. 
Bulletins  of  American  Paleonlologv  5(29-30):  165-459 
1=  1-2.52,  1-43],  pis.  27-65  |=  1-.39]. 

Morch.  ().  A.  L.   18.53.  Ciatalogus  conch\liornm  (juai>  reliquit 

D.  Alphonso  dWguirra  &  Gadea  Comes  de  Yoldi \ol. 

2,  Acephala,  Annulata,  Cirri])edia,  l'xliinodirmal;i.  Klcm. 
Hafniae.  |iv]  -f  76  pp. 

.Vlorch.  O.  .\  L.  I860  [18.59-1861].  Beitrage  znr  Molliiskcn 
fauna  f  A-ntral-Amerika's,  Malako/.oologische  Blatter  6(4): 
102-126  :1S59'.  7(2):  66-96,  (3):  97-106,  (4):  170-192 
(1860;,  (5i:  19.3-213  (lS61i. 

Olsson  A.  A.  1922  The  .Miocene  of  northern  Costa  Ric:L  uilh 
notes  on  its  general  siratigraphic  relations.   Bulk'tins  ot 


American  Paleontology-  9(.39):  1-309  [=  17,3-481  ].  pis.  4- 
35  pis.  [=  1-321. 

Olsson,  .'\,  A.  1961  Mdllusks  of  the  tropical  eastern  Pacific 
particularlv  from  the  southern  half  of  the  Panamic-Pacific 
lauual  province  (Panama  to  Peru).  Panamic-Pacific  Pele- 
cvpoda.  Paleontological  Research  Institution.  Ithaca.  574 
pp.,  86  pis, 

Pilsbiy,  H,  A.  and  H.  N.  Lowe.  1932.  West  Mexican  and  Cen- 
tral .\mericaua  mollusks  collected  bv  N.  N.  Lowe,  1929- 
31.  Proceedings  of  the  Aca(k'm\  ol  Natural  Sciences  of 
Pliiladelphia84:  33-144,  pis.  1-17. 

Pilsbn,  H.  A,  and  A,  A.  Olsson.  1941,  A  Pliocene  fauna  from 
western  Ecuador  Proceedings  of  tlie  Academy  of  NaturiU 
Sciences  of  Philadelphia  93  [for  1941]:  1-79,  pis.  1-19. 

Reeve,  L.  A.  1857,  Monogra]ili  of  the  genus  Siiiiij,iiin(>laiiiL  L. 
A.  Reeve,  ed.,  Couchologia  iconica;  or  illustrations  of  the 
shells  of  molluscous  animals  10:  1  pi. 

Rehder,  H,  A.  1967.  N'alid  zoological  names  of  the  Portland 
catalogue.  Proceedings  of  the  United  States  National  .Mu- 
seum l21  (.3579):  5rpp. 

Rckling,  P,  F,  1798.  Museum  Boltenianum  .  .  .:  pars  secunda 
contiens  conchvha  sive  testacea  uni\aKia.  bivalvia  &  mul- 
tivalva.  Trappii,  Hamburg,  \ii  +  109  pp. 

Rcimer,  H,  C,  1872  [1841-1873],  Die  Familie  der  Telhnus- 
cheln.  Telhuidae.  Svstematisches  Conchvlien-Cabiuet  von 
Martini  und  C^henmitz  10(4):  292  pp..  52  pis.  Gauer  and 
Raspe,  Niirnberg  [pis.  1-3,  first  issue,  1841.  bv  H.  C.  Kiis- 
ter;  pp.  1-.32,  pis.  4-6,  8,  10,  16,  1870:  pp.  3.3-176,  pis, 
7,  9,  11-15,  17-33,  1871:  pp,  177-2.56,  pis,  .34-38,  40,  .52, 
1872;  pp.  257-291,  pi.  ,39,  1873]. 

Sa\,  T.  1822.  .\n  account  of  some  of  the  marine  shells  of  the 
United  States,  Joimnil  of  the  .-^cadcniv  of  Natural  Sciences 
of  Philadelphia  2(2):  221-24S,  257-276,  302-325, 

Scliuiiuicher  C,  F.  1817.  Essai  dun  nouvean  svsteme  des  h:ib- 
itations  de  vers  testaces.  Sclmlfz.  Copenhagen,  |iv[  +  287 
pp..  22  pis. 

Spengler  L.  1794.  Noiere  bestemmelse  og  udvidelse  al  det 
Linneiske  gmnis  Soleii.  Skrivter  af  Naturhistorie  Sclskabet 
3i2i:  81-114. 

Spengler  L.  1798.  Over  det  toskallede  slaegt  telliuenie.  Skriv- 
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Tnon.  (;.  W..  |r  1869.  Catalogue  of  the  famiK  Tellinidae. 
Amrric.ui  journal  of  Conchologv  4(5)  [.\ppendix[:  72-126. 

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Willan,  R.  (.].  1993.  Taxouomit-  revision  nl  the  l:nniK  Psam- 
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Zeahmd  region.  Records  ol  the  \nstr:ili.ni  Museum,  Snp- 
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Wiicidrm'j;,  W.  P.  1982.  Geologv  and  paleoutologv  ol  ('im.il 
Zone  and  adjoining  parts  ol  Panama.  l!)escription  ol  I'er- 
tiarv  mollusks  (pelecvpods:  Propeannissiid;ie  to  Cnspuki- 
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to  tjasliopods:  cephalo])ods).  United  States  Geological 
SuiM'v  Professional  Paper  306F:  iv  +  .541-759,  pis.  8.3- 
124. 


THE  NAUTILUS  1  lfi(  1  ):13-24.  2{)()2 


Pam-  13 


The  unatom\'  and  functional  nioiphol{)g\'  of  Tivcia  vcnitricosa 
(Gray,  1838)  (Bi\al\1a:  Veneridae] 


Waller  Narehi 
Fahio  cli  Dario 

DcpurUiiiR'iito  (.If  Zoologia 
liistituto  de  Biociencias 
Uni\er.sidade  de  Sao  Paiiln 
BHAZIL 
wiiaifliiCS'usp.br 
lddario(S'ib. usp.br 


ABSTRACT 

A  tlctailcd  sIikK  oI  slicll.  organs  ol  tlir  iiiaiillc  ia\il\,  siplioiis, 
and  digcstixe  tract  of  TivcJtt  n-iithco.s/i  is  pre.st'ntcd,  and  lol- 
lowed  bv  conipari,sons  with  other  .specie.s  of  Tivrla.  Tiiclii  i  i-ii- 
tricosd  occurs  from  Espi'rito  Santo  State  in  Brazil  to  La  Palonia 
in  Urugnav.  Indi\idu;ds  li\e  in  .sand\  bottoms  (■\p<ised  to 
poiniding  .surf,  where  usualK-  a  large  aiiionnt  ol  nialrnal  in 
suspension  is  present.  The  anatoniv  and  Innctional  nioiphologv 
of  T.  ventricosa  are  compared  with  those  ol  congeneric  species 
occurring  oH  the  coast  of  southeastern  [Brazil.  The  si[ihons  ol 
T  vcntricosii  are  of  t\pe  B  of  Yonge.  the  ctenidia  ol  t\]ic  C.i  1 1 
of  .\tkins,  the  relation  between  the  labial  pal]is  and  the  etenitlia 
belong  to  categoiT  II  of  Stasek,  and  the  stomach  to  t\pe  \'  ol 
Pnrclion. 


INTRODUCTION 

Tlic  \  I'licroidca  is  a  large  superfaiiiiK  ol  liixaKcs  lliat 
lias  rachatt'cl  hroailK  worldwide,  t\picall\  into  soil,  iisn- 
all\'  intertidal  sediments.  The  genus  Tivcia  Link,  ISIIT. 
occurs  on  the  .Atlantic  and  Pacific  coasts  of  the  .American 
continent  (Abbott,  1974;  Coan  et  ah,  2(K)()). 

Isolated  references  to  Tiichi  iciifhcoMi  [('•vd\.  bSo.S) 
are  foimcl  mainlv  in  regional  checklists  (Lange  de  .\Ior- 
retes,  1949;  Gofterje.  1950;  Bncknp.  1957;  Hios.  197(1, 
1975.  19S5,  1994),  hut  tliere  are  no  icports  ou  ibe  lune- 
tioiud  inoipholog\-  of  the  species.  I'iicld  iciiliici'Mi  oc- 
curs Iroui  southeastern  Brazil  to  Urugua\  iHios,  19941 
.Along  the  Brazilian  littoral,  the  species  is  recorded  in 
the  states  of;  Espi'rito  Santo  (Rios,  1970),  Rio  cle  Janeiro 
(Rios,  1970),  Sao  Paulo  (Rios,  1970i,  Parana  (Cofferje, 
1950;  Rios,  1970)  and  Rio  Grande  do  Sul  (Bncknp.  1957; 
Rios,  1970,  1994).  The  species  was  rec'orded  in  Itios 
(1970,  1975.  1985,  1994)  from  the  littonil  of  Uruguay 
and  Broggi  (1970)  collected  hissil  representali\es  ol  the 
species  from  the  Qiierandino  FoiiiKitidu,  Uruguax.  and 
described  the  distribution  of  the  fixing  animal  Irom  San- 
tos (Bnizil)  to  La  Paloma,  Rocha  (Uruguayi.  Tiichi  n-ii- 


//■;V(<sy;  was  not  recorded  li\  (.'arcelles  (1944'  and  (.'ar- 
celles  and  Williamson  (I95|i  Irom  |Ik>  coast  ol  Argen- 
tina. 

.Although  not  sold  ui  llie  coinmereial  seafood  markets, 
the  species  is  an  import;nit  food  item  for  coastal  po])u- 
lations,  tourists,  and  campers  on  the  coast  of  Sfio  Paulo 
State,  Brazil,  The  main  goal  of  the  present  paper  is  to 
proxide  basic  inlonuation  on  the  ,matom\  and  some  as- 
pects ol  the  lii<ilog\   ol   the  species. 

.M.ATEfilALS  ANf)  MI"ni()l)S 

Li\ing  specimens  of  T  niilricd.sri  (Figure  1)  weri'  I'ol- 
lected  on  the  intertidal  region  of  beaclies  in  the  localities 
of  Sao  Mcenle.  Bertioga.  and  Ubatuba.  all  on  the  coast 
of  Sao  PiLulo.  Bnizil.  Specimens  were  lound  buried  at 
depths  of  appro\iiuatel\  5  cm  in  sand  oi'  nuiddy-sand 
substrates,  in  areas  with  strong  dist)n-|)anc<'  of  bottom 
deposits,  wbicli  end  up  suspended  in  large  amounts  in 
tlu'  wati'r  column.  Specimens  wert'  collected  during  low 
title,  when  lhe\  e<inld  be  detec-ted,  at  the  surface  of  the 
substrate,  b\  ,i  little  circular  mound  deposited  around 
the  tips  of  liie  siphons.  Some  of  the  study  animals  were 
kept  aliM'  np  lo   I  months. 

Drawings  wi^re  m;ide  from  relaxed  and  pri-seneil 
specimens.  .Vhignesium  sul|)liate  was  used  as  a  relii-xing 
agent,  ('iliarv  currents  wei'e  studied  with  the  use  of 
Carborundum  (F),  carmine  suspensions,  or  suspensions 
of  .A(|uadag.  Organs  and  tissues  were  fi.xed  in  Bouin's 
fluid,  and  sections  (i-S  |jLm  in  thickness  stainetl  with 
Ehrlich's  baematowlin  .uid  eosin.  .\z;m.  and  .Shillon's 
triple  stain. 

N'oiichcr  specimens  are  dc|)osiled  in  the  Museu  (ic 
/,oologi;i.  Uni\ersid;ule  de  Sao  Paulo.  M/Sl'  3294S. 
.32949. 

RESULTS 

.Shell  (Fijiiires  2—4):  4"bc  shell  of  T  iciitricosti  is 
e(|ui\;iKc.  ei juikitciiil.  trigonal,  with  subcentral  nndxi- 


Pai^c  1  t 


THE  NAUTILUS,  \o\.  116.  X( 


r<'>D^  or^o  0  , 


■'O-Q  < 


/fo  .0*-J.c'?C>>-r 


0 


^ 


■i  O    oo  ,   O^O  oCi,    ,,>^^0    Or,  9  0° 


C^'^oCP^, 


15,  Po 


g  O  O  0,2    O/ 
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:o^^  ■ 


0 o(>?<^"^  ao'5^&«^^o^:V^d 0, 


^  -  q;?^6<^^>^^ ;  ^^^^^^o  c>  V  0 


<5b 

0- 


Figure  1.     Tivchi  icnlrirosa.  Aiiiiiuil  iii  iiatmal  li.iiiital:  cvtci- 

rial  \ie\v  Iroiii  the  left  side.  Arrows  show  the  direeti I  ihc 

iiRiirrent  and  exeurrent  currents.  .Scale  line  =  2  cm. 


lies  and  beaks  located  close  to  eaeli  oilier  Tlie  external 
surface  is  smootli,  oxcrlaid  1)\  a  tliiek.  <j;loss\  [lerios- 
traenm.  which  gix'es  a  varnisiiccl  appearance  to  the  shell 
vaKcs.  The  external  color  is  cream  with  a  xariahle  pat- 
tern of  radial  or  zig/a<4  hrown  streaks  (Fimire  2).  Sonie 
specimens  show  alternating  ime(|nal  ra\s  of  straw  and 
chestnut  color,  with  the  rav  color  on  llie  ri'^lit  \al\e 
olteii  more  \ivi,l  than  that  on  the  left.  The  interior  ami 
respeeti\e  Iree  inartjiu  ot'tiie  shell  haxc  a  smooth  whiti' 
surlace.  The  pallial  sinus  is  short,  ohiiijue.  U-siiaped. 
The  .scar  of  the  anterior  adductor  imisele  is  o\al  and 
more  impressed  than  the  posterior  one.  which  is  almost 
circular  in  outline  (Figure  3).  The  hinge  is  strong,  with 
3  weli-dexcloped  cardinal  teeth  in  each  \alve  (Figure 
4).  The  uiedi.m  toed,  is  n.irrower.  hiadedike:  the  an- 
terior and  posterior  sets  are  hilid.  with  the  posterior 
one  coar.seK  .serrated.  Thi^  knoh-like  anti'iior  lateral 
tooth  on  the  left  \al\e  fits  into  a  deep  dejjression  on 
the  riglit  \ai\e.  Along  tlie  posterior  dorsal  margin  of 


the  right  \al\e  there  is  a  large  grotne  and  a  smaller 
groove  is  present  on  the  anterior  dorsal  margin  of  the 
teft  \al\e.  Hie  shell  of  the  largest  studied  specimen 
measured  7.U  cm  in  length.  7.4  em  in  height  and  5.2 
cm  in  width. 

Siphons  (Figures  5,  6):  Thi'  siphons  are  short  and 
fused  at  the  base.  They  are  formed  bv  fusion  of  the  inner 
fold  with  the  internal  surface  of  the  middle  fold  of  the 
mantle  margin.  The  aperture  of  the  incurrent  siphon  is 
surroimded  bv  branched  tentacles,  and  that  of  the  e\- 
iiirrent  siphon  has  a  tapered  siphonal  membrane  with 
simple  tentacles  at  the  basal  region. 

The  incurrent  siphon  (Figures  5,  6)  has  about  200 
tiiitacles  deployed  in  5  sets  surrounding  the  aperture. 
The  tentacles  are  directed  toward  the  aperture  when  the 
animal  is  pumping  water  in.  which  allows  for  sieve-like 
trapping  of  particles.  The  most  e\tern;il  set  consists  of 
small,  digitiform  tentacles,  some  of  which  displax"  a  bi- 
furcated tip.  The  four  internal  sets  consist  of  branched 
tentacles  that  decrease  in  ninnber  and  increase  in  length 
in  each  succeeding  set  toward  the  internal  region  of  the 
siphonal  opening.  A  U-shaped  pigmented  dark  band  is 
present  at  the  base  of  the  group  of  three  most  internal 
sets  of  tentacles.  The  incurrent  siphon  is  \"er\'  sensitiw 
to  touch  and  \ibrations  and  witlidiaws  in  part  oi'  com- 
pk'teK  upon  stimulation. 

The  excurrent  siphonal  aperture  (Figure  fi)  is  smaller 
than  the  incurrent  siphonal  aperture,  .\bout  4S  simple, 
external  tentacles  of  niilk"x-white  color  surround  the  base 
of  the  siphonal  uieiiibrain\  These  tentacles  are  pig- 
mented at  their  bases. 

Siphonal  Menihranes  (Figure  7):  .\  pair  of  siphonal 
membranes  is  prest-nt  at  tlu'  basal  opi^ning  of  the  incur- 
rent siphon;  these  membranes  eoiitribute  to  regulate  the 
water  flow  into  the  pallial  ca\it\.  Two  other  additional 
si|)honal  folds  are  present  and  unitetl  at  the  dorsal  side 
of  tlu'  incurrent  siphonal  opening;  thev  eontimu'  \en- 
tialK  and  in  anterior  ihrection  as  expansions  of  each 
iiiaiitle  lobe. 

Organs  of  ihe  Mantle  Canity  (Figures  <S,  9):      He- 

mo\al  of  the  left  shell  \al\c  and  respectixe  mantle  lobe 
exposes  the  organs  in  the  iiumtle  c;i\it\-  (Figure  8).  The 
ctenidia  ;n"e  dorsalK  loe;ite(l.  The  inner  deiiiibrancli  is 
kirger  than  the  onlei  one;  this  latter  bears  .i  snpraxi;il 
extension,  Tlie  imiei  iiumlle  siirf;ice  is  coxered  with 
slioit  cilia,  except  in  a  narrow  border  on  its  free  edges. 
The  mantle  edges  are  fusi'd  for  ;i  short  extension  ;mte- 
ii<ir  to  die  iiK'iirreiil  siphon,  1  )ors,il!\  lo  these  fused  \en- 
Ir.il  m;i|ij;ins.  the  lii'j,lil\  mobile  ;idilitioiial  siplimial  iiiein- 
bi;iiies  delimit  a  eaii.il,  Ihe  Iree  edge  of  the  iiiaiille  li.is 
4  folds  and  a  large  number  of  ramified  tentacles,  (hose 
ill  the  proximal  and  distal  regions  ol  (he  pedal  gape  di- 
\uled  into  seeondai'X  and  (erli,ir\  biaiiehes,  'I'hese  short, 
simple  tentacles  uia\  ;ilso  be  ;ibsent  in  the  niedi;m  region 
of  the  pedal  gape.  When  tlie  ;miiiial  is  resting,  the  ten- 
tacles on  the  ielt  side  .liteniale  with  those  on  ihe  right 
side,  so  til, it  lhe\  prevent  sand  gr;iins  Irom  entering  IIk' 


W.  Naalii  and  F.  di  Darici.  2002 


Page  15 


Figure  2.      Tiicla  rciitrirnsa-  Extfiiial  xirw  <A  slicll  \aKcs  sIhiwIih,;  \ariali(iii  in  cdliir  pallcrii.  Scale  line  -  2  cm. 


mande  ca\it\-.  When  the  foot  is  protruded,  tlie  tentacles 
touch  all  its  sides,  forming  a  protecti\e  latticedike  stnic- 
ture  around  it. 

Cilian  currents  on  a  wide  area  of  the  mantle  surface 
are  ventralK-  directed  toward  txvo  main  rejection  tracts, 
one  on  each  \entral  margin  of  the  mantle  lobes.  The 
main  rejection  tracts  drixe  and  accumulate  psentlcjiaeces 


into  the  xcntral  canal;  this  latter  protects  undesirable 
material  from  being  washed  back  into  the  mantle  i-ham- 
ber  (Figure  9).  Upon  accnmnlalion  of  a  gixcn  amount 
ol  pseudotaeces.  the  adductor  muscles  snddeuK  contract 
to  pusli  watci"  out  ol  tin'  mantle  ca\it\  causing  the  si- 
|ihonal  membranes  to  open  and  allowing  pseudolaeces 
to  be  eliminated  through  the  incurrent  siphon. 


Page  Ifi 


THE  NAUTILUS.  Vol.  116.  No.  1 


Figure  3.  Tiiclti  nittiicusd.  Iiitriiial  \U'\\  (if  tlie  riij;lit  shell 
\;iKe,  .slumiiig  tlic  pallia!  .siiuis  anil  the  scars  (il  the  adductor 
muscles;  an.  auterior  adihutor  nuiscle  scar;  /),  pallia!  liue;  pa. 
posterior  adductor  uuiscle  scar;  ps.  palli;il  sinus.  Scale  liue  = 
3  cm. 


Mu.sculature:  The  anterior  adductor  mu.scle  is  larger 
liiiui  ll]c  postciior  one.  The  anterior  pedal  retractor 
nuiseles  are  inserted  into  the  shell  \al\e  posterior  to  and 
at  a  short  distance  from  the  anterior  adductor  nuisele. 
Some  ol  its  fibers  spread  superficially  into  tlie  anterior 
and  dorsal  region  of  the  foot,  while  most  spread  in  pos- 
terior direction:  thest'  latter  have  tlu^r  t'xtremities  in  the 
foot  intcrnallv  to  the  fibers  coining  IVoni  the  posterior 
pedal  retractor  muscle.  DistalK,  within  tlie  foot,  the  fi- 
bers from  both  sides  join  together. 

The  posterior  ped;il  retractor  muscles  are  inserted  in 


Figure  4.  Tiuhi  i,;,fn,,n,i  luterual  views  of  the  shell 
valves,  details  of  teelli  Top.  left  \aKc;  bottom,  right  \aKc 
Scale  line  =   I  cm 


Figure  5.  I'iiflii  loitiiriKd  FulK  exleiided  incuneut  siphon 
projecting  out  ol  the  shell  \al\es,  showing  the  intricate  dispo- 
sition of  the  branched  tentacles.  Scale  line  =  0.2  cm. 


the  shell  \al\e  in  .uitciior  direction  and  dorsalK  to  the 
posterior  ;iddnctiii"  muscle.  The  letractor  nuKScles  on 
each  side  unite  inuler  the  pericardiiun,  where  bundles 
of  fibers  Imm  the  light  nnrscle  cross  with  those  from  the 
left  one.  Where  the  nuiscK's  enter  the  foot,  their  bundles 
once  more  dixcige  and  their  fibers  radiate  in  anterior 
directinn  ;uiil  xenli.ilK   nitn  ihc  loot. 

The  pedal  retnictor  umscles  constituti"  the  extrinsic 
muscuhiture  ol  tin'  loot.  In  ;id(litiou  to  the  extrinsic  nnis- 
culalurc.  ;i  \;n-\ing  amimnl  ol  intrinsic  nnisciilature  is 
also  picscul  in  the  pidximal  ;md  dist;il  portion  ol  (he 
loot.  Innnedi.ilrix  liclciw  the  epithchnni  iil  the  \iscer.ii 
mass  in  the  pi()\im;il  p;ut  ol  the  loot  there  are  circuhu' 
muscle  strands  lh;it  run  in  ;mtcro-posterior  direction  and 
completeK  suiT'ound  llie  \isccr.il  in;iss.  The  distal  Iree 
[)art  ol  the  hiot  is  patki'd  with  bundles  ol  fibers  running 
transversiilK,  ihe  same  occiuiing  in  llie  \iscci;[l  nuiss, 
wlieic  transverse  libel's  ;u"e  less  nnnicnins. 

Foot:  The  loot  is  hirge.  wcdge-slKi|)ed.  ;uid  fileralK 
comjiressed,  which  laiilitales  burrowing  in  I'ehitivcK  soft 
substiiituni,  where  il  di<fs  with  rlixllunie  movenicnfs.  .\ 


\\\  Xarclii  and  F.  (]i  Dario.  2002 


l'ae<*  r 


sm 


Figure  7.  In  i  In  i  nifriidsd  Internal  \  icu  iil  llir  ci[)('i]lii<j;s  ol 
the  ineiirreiit  ami  r\riinent  siphons  in  the  mantle  ea\it\':  f/.v. 
additional  siphonal  nienihrane:  in.  inenrrenl  siphon;  ins,  ineur- 
lerit  siphonal  nienihrane;  c.v.  excnrrcnt  siphon;  ]>ri,  posterior 
atldnetor  nnisele;  shl  siplioiuil  nnisele.  Se;ile  line  =  1)5  em. 


Figure  6.  Tiiclii  vcntiicosn.  ExttMuletl  sipli(ais  seen  Irom 
al)o\e.  Top,  diaijrani  emphasizini;  disposition  ol  tiii'  5  sets  ol 
tentacles  of  the  inenrrent  siphon;  c.v.  exciirrent  siphon;  in.  in- 
eurrent  siphon;  I,  II  III.  I\'.  and  \'.  sets  oi  tentacles.  Scale  line 
=  0.5  cm;  liottoni,  detail  of  the  openiini;  ol  the  exenrrent  si- 
phon showing  the  openinii  nK'inliniiie  and  tentacles.  Sc;ili'  line 
=  0.2  cm. 


its.'ir 


e(iiii| 


)letfl\ 


specimen  ol  4.S  (.■iii  shell  lenu;tli  hiiried  it 
in  7  minutes. 

.Specimens  h\e  compIeleK  liuried  at  a  depth  ol  ap- 
pro.ximatelx"  5  em:  olten  specimens  can  lu'  limnd  with 
tlie  shell  pointiii'j;  out  nl  the  substratum. 

Ctenidia  (Figures  8,  10-12):  The  shape  ol  the  cte- 
nidiii  and  the  current  patterns  on  them  are  illusti;ited  in 
Figures  8  and  10.  Each  ctenidium  is  Formed  l)\  t\ui  de- 
mihranchs,  the  inner  demihranch  hi<j;lier  than  the  outer 
one,  particularK  in  anterior  direction,  and  i:;ioo\ed  iilon'j; 
its  free  margin. 

Tlie  lamellae  olliotli  dtanihranchs  are  plicate  ;md  liet- 
erorhalxlie  (Figure  1 1 ).  Plicae  are  present  alonii  the  larg- 
er part  of  both  lamellae,  ranging  from  a  mininmm  ol  29 


to  a  maxiiinmi  ol  (iU  in  the  outt-i'  and  imiei'  demibr;uiehs. 
The  filaments  (Figure  12 1  bear  lateio-trontal  (fi  |Ji.m 
long)  and  frontal  cilia  (10  |jLni  long),  the  latter  replaced 
bx'  increasingk-  longer  terminal  cilia  (uj-)  to  40  |xm  long) 
in  the  distal  free  edge  of  the  filament.  Throughout  the 
lateral  sides  of  the  filaments  lateral  cilia  produce  a  pow- 
erfid  respiratoiA  and  feeding  current. 

On  tlie  ascending  lamella  ol  the  outer  demihiMiieh. 
downward  eili;ir\  eiiirents  were  obsened  on  ;ill  filaments 
of  the  eicsls  iiiid  lioiighs  of  the  plicae,  (.'ollected  mate- 
lial  is  iiKiiiiK  eon\e\{'d  around  the  free  margin  of  the 
outer  ilemibraneh  onto  its  descending  lamelhie  where 
frontal  cilia  e;iri-\  |i;irticles  onto  an  acceptance,  month- 
directed  current  on  the  etenidi;il  .i\es. 

There  is  no  food  groo\e  along  the  free  edge  ol  the 
outer  demibranch,  but  a  group  of  terminal  cilia  along 
the  oiilei'  fiee  delleets  large  particles  in  anterior  direc- 
tion. These  p.iilicles  are  carried  lor  a  short  distance 
;iloiig  the  free  iiKUgin.  and  then  tr;uislerred  to  the  outer 
surface  of  the  iiiiiiM-  demibranch.  On  the  Irontal  region 
of  the  outer  deinibrancli  tliere  are  \t'n  large  cilia  that 
cle;m  the  ctenidium  b\  lenuning  partit'les  as  large  ;is 
sand  ^niins, 

'file  lroiit;il  cilian  eiirri'iits  on  both  l.miell.ie  ol  the 
inner  demibranch  are  almost  alwa\s  directed  toward  the 
marginal  grooxc,  e\ce]ition  made  to  thv  proximal  region 
of  the  descending  Limellae,  where  cilia  be;il  in  dors;il 
direction  to  eoii\e\  material  to  the  aecept;nice  tract 
aloii'j,  the  elellidium  ;ixis. 

Labial  Palps  (I'igiiri's  13,  14):  'fhe  l.ibi.il  palps  ari' 
flat,  triangular,  with  the  inner  faces  tleepK  plicate  and 
the  outer  faces  smooth  (Figure  13).  The  \cntral  tips  ol 


F;l<TL'  IS 


THE  NAUTILUS.  \ol.  116,  Xo.  1 


U 


ex 


Figiirt'  S.  Titi'la  icnlrirosii.  Oiwaus  of  thf  iiiHiitlc  ( a\it\  \iiwr(l  Ikhii  llir  hit  side  alter  R'liiox.il  (iltlic  Iclt  shell  \.i\w  .mil  Ictt  in. mile 
lobe;  (III.  iuiteiior  adductor  iiinscle:  ti.s.  atklitional  siplioiial  iiieiiilnane;  ex.  cxcuiTenl  siphon;  /.  loot;  id.  inner  deniihianeli;  ilp.  inner  labi;il 
palp;  ill.  incniTcnt  siphon:  m.  mantle  edije;  oil.  outer  deniilinmeh;  nip.  outer  labial  palp;  ji/i.  posterior  ;i(lductor  ninscle;  sr/c.  supra-axial 
extension  ol  the  outer  ileniil)r;uieli:  //.  iiuilio;  i,  \entiiele.  Anows  iudiciite  the  direetiou  ol  the  eurrents.  Se.ile  line  =  2  em. 


tilt'  most  anterior  filaments  of  the  inner  deniiliniiuli  ;ire 
inserted  into  and  Insed  Id  the  (list. [I  m-.il  ;j,iii()\e  ni  ihe 
palps.  On  tiie  antero-doisal  ri'tjion  nl  the  external  snr- 
faees  ot  tlie  palps,  ei!iar\-  enrrents  move  particles  aionnd 
tlie  dorsal  margin  to  the  internal  sinfaees.  The  following 
ciliai"\-  currents  on  the  |)alps  (I'"ignre  I  li  weic  I'eeog- 
nized: 

1.  Acceptance  currents  (a)  that  condnct  parti(  les  to- 
ward the  nionth  I'roni  one  crest  of  the  plic-ae  to  liie 
next  ib:  dois.illv  directed  on  the  ahorai  surface  of  e;ic'h 
plica,  and  eL  on  the  lateral  region  of  the  folds,  a  vvu- 
trally  directed  current  is  present  on  the  crest  of  eacli 
ridi^e. 

2,  Hej(eiii,M  lurrenls  id),  on  the  lloor  of  troughs  he- 
txveen  .idj.ieeni  plleiie  :ui(l  (e)  on  the  lower  half  of  the 
sides  oi  til'-  plii  .1  ^\her(■  particles  are  driven  onto  the 
smooth  \<(iir,il  eil'^e  .)f  ihe  p.ilps.  Here,  a  strong  rejec- 
tion ciM-renI  e. ,uiir,els  m;ili'ri;il  towards  the  free  tip  of 
the  palp. 

Mn.scnlar,  as  •ai  I!  :is  i  ilian  ;icti\itx-  nmst  he  taken  into 
account  when  tin     lelion  of  the  laliial  pali^s  in  sortimj; 


lood  particles  is  considered.  The  palps  ol  T.  vcninrosa 
;U('  i"<'hiti\el\  ;ieli\{'.  The  plicae  on  the  iiuiei"  snii.iee  nia\ 
he  erected  or  \mi.\  Hal.  ;ni(l  the  p.ilps  uui\  lie  contractetl 
and  twisted  into  ;i  spii;il  shape,  h I'i n ti;i n 'j;  th<'  smooth  xcn- 
ti;il  ed<j;e  .uul  icspeelixc  lejet-lion  c'nrrent  in  contael  with 
the  ridged  sui'hice. 

When  siu;ill  amounts  ol  [Kulieles  .tie  presented  to  the 
p;ilps.  little  sciiting  (;ikes  phiee.  The  lidges  are  I'elaxed 
and  oM'rlap.  Ivxposed  to  the  ;iccept;mce  currents,  the 
majoritx  ol  [)articles  are  c.uried  rapidK'  toward  the 
month. 

Laige  amounts  of  iiarticles  on  the  palj)s  induce  dil- 
lereut  deijrees  of  nmsenlar  acti\it\;  which  results  in  o[i- 
tinud  soiling  ellieiencw  I'.xcess  ui;itei"i;il  e;uises  ,i  nms- 
enlar reaction  In  contraction  ol  the  organs  into  ;i  s])iral 
shape  and  most  material  to  he  ri>jeete(l. 

.Suliuiilted  to  moderate  amounts  ol  uiateri;il.  the  \ar- 
ions  sorting  anil  icjectiou  cmicnts  on  the  plie;ie  are  ex- 
posed. Large  particles  fiilling  into  the  troughs  ol  the 
gro()\es  are  rejected,  while  ihe  sm;dler  ones  ;ii-e  trans- 


W.  Narelii  and  F.  di  Dario,  2002 


Faw  19 


sae 


Figure  9.  Tinlii  icntricosti  Iiiiu-r  siirlaci'  of  the  ri^lit  inaiitlt' 
lobe  alter  reni()\;il  nf  the  lelt  shell  \al\e  and  part  i)f  tiie  left 
mantle  lobe  sjiowing  eiiian-  cieansiiis;  currents  and  additional 
siphonal  membranes.  Arrows  indicate  the  direction  ol  the  cur- 
rents. Scale  line  =  2  cm. 


porti'il  In  the  sortiiiLi;  currents:  oiiK  pari  (il  the  original 
material  reatiies  tlie  oral  i^rodxe  hetween  tlic  palps. 

Alimeiitai-\  Canal  (Figures  15-lfi):  A  short  isoph- 
ai;us  opens  nito  the  anterior  wall  ol  the  stomach.  The 
cornhined  st^le  sac  and  intestine  opens  into  the  poster- 
oxt'iitral  region  ol  the  stomaeli.  and  passes  l)ack"\\ard  and 
downward  as  a  wider  tulie  into  the  loot.  The  mid'j;nt 
continues  Ironi  the  distal  end  ol  that  widei'  tube,  coils 
tisihtk  about  fixe  times  on  the  xt^ntral  side  ol  the  stom- 
ach, then  passes  downwartl  and  back\\ard  to  ascend  as 
die  hintl<j;ut  in  posterior  direction  to  the  st\le  sac.  TIr' 
hindL:;ut  passes  through  the  pericartliuni,  wheic  it  is  sur- 
rcjuudcd  1)\  the  ventricle  antl  posterior  aoitie  bulb,  and 
lings  dorsalK'  the  posterior  adduitoi'  muscle,  to  end  in 
the  anal  papilla. 

The  stomach  (Figure  16)  has  its  esophageal  opening 
defined  In  a  transverse  rim.  The  minor  t\phlosole  ends 
on  the  right  side,  close  to  the  orifice  of  thi^  combined 
st\le  sac  and  intestine  in  the  floor  ol  the  stomach.  Tlie 
major  tsphlosole.  associated  with  the  intestinal  gjoove, 
penetrates  the  stomach  and  passes  loiAxaiii.  then  down- 
ward into  the  right  caecum.  Tlie  riglit  caecum  receives 
six  ducts  Irom  the  digestive  diverticula.  Emerging  iroui 
the  right  caecum,  the  major  tvphlosole  passes  to  the  lelt 
across  the  floor  of  the  stomach,  forming  a  flap  that  pro- 
jects in  dorsal  direction  toward  the  esophageal  orifice. 
The  major  tvplilosole  then  is  directed  deepK  into  tlii' 
lelt  caecum,  returns  and  ends  on  the  caecum  opening. 
The  left  caecum  receives  4  ducts  Irom  the  digestive  di- 
vi'rticula.  The  origin  of  the  intestinal  gi'oovc  lies  on  the 
lelt  anterior  floor  of  the  stomach,  close  to  the  opening 
ol  the  left  caecum. 

CiliaiT  currents   in    the   lelt   caecum   diicet    jiaitick's 


alod 


a  I  id 


Figure  10.  Tii  i-hi  i  citlntOMt  l)i.ii;raimnalic  vertical  section 
through  tlie  cteiiidia  m  tin-  stvle  ol  .\tkins  ( 19.37b)  showing  the 
direction  ol  particle  IIoh  bv  llic  Irontal  cilia:  alid.  ;LScending 
lamellae  ol  the  inner  demihnincli:  alod.  ascending  lamellae  of 
the  outer  demibrancli:  (Uid.  descending  lamellae  o(  the  inner 
deniibranch:  dlod.  desciMiding  lamellae  ol  the  outer  demi- 
brancli: g.  marginal  groove  ol  the  inner  demibrancli:  sar.  supra- 
avial  extension  ol  tlu'  outer  deniibraiuii.  .Arrows  indicate  the 
direction  of  the  cilian  currents. 


:iw:iv  Irom  the  orifices  ol  the  ducts  earning  them  toward 
the  intestinal  groove. 

A  group  ol  5  ducts  lioin  the  digestive  diverticula  en- 
ters the  left  pouch.  A  soiling  area  King  on  the  floor  of 
this  pouch  prolongs  onto  the  right  side  ol  the  stomach, 
where  il  exjiands  and  lornis  a  bi-aded  swelling.  Cilia  on 
this  sorting  area  beat  backward  along  the  grooves  and 
awav   Irom  the  openings  ol  tile  ducts. 

.\  wi'll-deveioped  dorsal  hood  projects  upwards  Irom 
the  rool  ol  the  stomach  ami  cunes  over  toward  the  left 
side.  .\  sorting  ari'a  on  its  rool  and  anterior  wall  prolongs 
over  tlie  esojihageal  opening,  then  on  tlie  right  wail  of 
the  stomach  to  finisii  posterior  to  tiie  orifice  of  the  right 
cai.'cinii.  Material  rejected  Irom  this  sorting  area  is 
passed  into  a  rejection  tract,  wiiicli  arises  witliin  the  dor- 
sal hood  and  discliarges  into  the  intestinal  groove.  Tlie 
:interior  margin  ol  the  rejection  tract  is  fornu'd  hv  a  fold 
v\ilh  sevenii  small  ridges. 

'I'lie  gastric  shield  covers  an  extensive  area  on  tlii'  left 
wall  ol  the  stomach,  and  sends  a  flange  into  the  opening 
ol  die  dorsal  liood,  and  anotiier  into  tlic  lelt  poucii.  Vhv 


FiU'e  20 


THE  NAUTILUS,  Vol.  IIH.  N( 


%«^ 


d>ill§§ 


Fij^urc  11.  Tin'Ia  vcnlricDsa.  Tninvfisf  sixtioiis  cil  tlic  outi-r 
(l('iMii)iaiKli.  Top,  niL'clian  plicate,  lieterorhabdic  condition  of 
the  ctcnidiuni.  Scale  line  =  0.1  nun;  center,  detail  of  the  plicate 
deniihrancli.  Scale  line  =  ().2.t  nnn:  bottom,  di.stal  non-plicate 
condition  ol  (lie  onter  deiniliiancli.  Scale  line  =  0.1   nnn. 


^ 


l^l«, 


s — "' 


/    I   '   /    ^  I 


h\ 


Figure   12.     Til ,  / 
Ix'ft.  diagram  oftlH 
rij;Iit.  detail  of  two  li' 
current; /r.  froiilal  cili, 
re,,  larjie  frontal  cili;i;  /' 
direction  ol  tlie  ciliarv  cnrn 


ilialion  p;ittern  on  clenidia. 

'    '  snri;ice  ol  the  ctenidinni; 

•     llie  lat(Mal  cilia;  a.  oral 

'i.i,  lfi\  laterolrontiil  cilia; 

'la.  .Vrrow.s  in(lic;ite  the 

..  •    lini-  =  .50  (iin. 


Figure  13.  iiitld  imlricdsa.  Cilian  cnrreiits  on  the  labial 
palp.s.  Top,  external  .surface  of  the  labial  palp;  bottom,  intennil 
.sin'face  of  the  labial  palp;  din.  dorsal  mary;in:  (7/j,  inner  labial 
palp;  ()//j,  outer  labial  palp;  viii.  Nfntnil  margin.  Sc;de  line  =  3 
cm. 


ga.stric  sliicld  li:is  ;i  small  toiitli-likc  projection  ;it  tlic  an- 
terior eiul 

There  is  a  .sorting  area  on  the  anterior  wall  ol  the 
storn;ieh.  het\\t'en  the  aperture  of  the  esophagus  and  tlie 
tr;ins\-erse  section  ol  the  intestin;il  grooxe.  (Mlia  on  this 
area  beat  upuaril,  toward  the  esophagus.  This  sorting 
area  extends  to  the  right  caeeuni.  where  cilia  heat  along 
the  grooxes  from  rigiit  to  left.  ;uul  proh.iliK  eon\e\  p;u"- 
ticlt's  toward  tlie  dor.sal  hood. 

i:)is(;ussK)N 

Tilda  I'ciilricii.sii  is  restricted  to  sand\   heaches  direetiv 
exposed  to  wave  action.   The  species  is  no(  lound  in  shel- 


^^....i-i iuV-:y':-'i''Vi 


Figure  14.  Tiirlti  vciitricosd.  Diagnnnnwtic  representation 
ol  the  ciliinA  mecli;nnsnis  on  the  lolded  inner  snrhice  of  the 
l;ibi;il  palp,  sliouint;  the  \;irj(ins  cili;ir\  tnicts.  .\nlcri(ir  is  ;il  Icif 


W.  Narchi  and  F.  di  Daiio.  2002 


Pat'c 


Figure  15.  T'ncUi  icntrircsii  AliiiicntaiA  caiiaL  sirii  Ikhii 
tlic  lelt  side;  nil.  anterior  atkliictor  iiiiiscle;  up.  anal  papilla:  </(/. 
dii;esti\e  di\erticula:  ilh.  dorsal  hood;  c.  esoplianns  f.  foot;  /ii,'. 
liindgiit;  mo.  mouth;  pa.  posterior  adduetoi'  ninselr;  ;  .  ventri- 
cle: .s.v,  st)'le  sac;  .st.  stomach.  Scale  line  =  4  cm. 

tcrt'd  lia\s  or  other  en\ironin('nts  lacking  strong;  wave 
action.  Tiiis  sii<itie.sts  that  a  liiti;li  tleifree  of  :u'ration, 
made  possihk'  l)\  stroii'j;  niixiiiti  at  the  uater/air  iiitei- 
lace,  i.s  essential  to  tlie  species.  (loderje  (1950)  lound  T. 
vcntiico.'ia  on  open  seashores,  co-occurring  witli  the  \e- 
nerid  hWdWv  Aini/inti'i  piirpunitiis  (Lamarck,  ISIO).  Til- 
da rciitriciisa.  like  Tilda  stultonnn  (Mawe,  1S2.'5),  as 
tlescrilieil  h\  Wevmoudi  (1920,  1923),  is  exposed  to  con- 
stant sin-f  pounding,  whicli  its  thick  shell  is  ahle  to  with- 
stand without  cracking.  The  species  apparentk  thrixes 
in  what  might  he  c'onsiilered  as  an  inil;i\i)r:il)le  cnxiron- 
ment. 

The  large,  liea\A',  massi\e  shell  ol  7'  Kiiliicusa  ex- 
ceeds in  size  all  other  species  of  Tilda  occurring  oil  tlie 
Brazilian  coast.  The  largest  specimen  collected  during 
the  piesent  stndx'  (7.9  cm  in  li'ngth:  7.4  cm  in  lieigiit. 
and  5.2  cm  in  width)  exceeds  the  largest  one  registered 
by  Gofferje  (1950)  (3.5  cm  in  lengdi,  3.0  cm  in  height, 
and  0.9  in  widdi). 

Armstrong  (1965)  uiulertook  sonic  experiiiiciits  willi 
10  species  of'bi\ai\es,  including  Tilda  slullaniin.  .Study- 
ing the  position  of  the  animals  in  relation  to  die  w:i\c 
front  and  tlieir  lielia\ior  in  the  sulistratinii.  tli:it  .uitlicn 
found  that  the  direction  of  \va\e  action  is  related  to  the 
//(  sitit  orientation  of  that  species.  Such  behavior  was  not 
observed  in  tliis  stmK'  for  T.  icatriccsa.  nor  lor  7'.  inac- 
froidrs  (Bom,  177.S) '.studied  by  Narchi  tl972i.  .Ml  lol- 
lowing  references  to  T.  mactroidcs  are  from  Narchi 
(1972)  unless  othenvise  noted. 

The  siphons  of"  T!  iciithcosa  aw  ol  T\pe  15  of  Voiige 
(1948,  1982).  The  extremek  moliile  siphonal  membrauc 
of  the  excurrent  siphon  interferes  with  the  flux  of  water 
passing  through  the  ctcnidia:  when  opened  and  extend- 
ed outwaiil,  the  membrane  controls  and  directs  the  I'x- 
halant  current.  .As  observed  in  T  luactnudcs.  there  is  no 


leiitiKiilar  ring  loiined  In  the  iiicdian  numtk-  fold 
around  the  excurrent  :ind  incurrent  :ipi-rtiires  in  T.  icii- 
Irifiisa 

The  siplioiis  of  7'  iciilricdsa  are  similar  in  stmctiire 
and  shape  lo  diose  described  lor  T.  sluUoniiii  (W'ev- 
iiioiilh.  1920.  192.3)  antl  T  iiiaclroidc.s.  The  incurrent 
siphon  in  T.  icnlricosa  has  branched  tentacles  disposed 
in  5  sets  surrounding  the  a])ertine,  unlike  T  miHiroidvs 
where  tliev  are  disposed  in  3  sets.  Thi'  excurrent  open- 
ing of  i.  icnlricosa  has  about  48  simple  tentacles,  which 
are  darkK  jiigmented  at  tluir  l);ises,  wliile  in  T.  inactro- 
idc.s  the  excnrrenl  opening  teiitacli's  are  :ibout  20.  and 
niilkv  while  in  eoloi. 

Narchi  :iiid  I  .opes  (19981  recorded  for  tin-  hrst  time 
the  presence  of  the  additional  sijihonal  membranes  in  T 
icnlricosa.  In  this  sjiecies,  the  tx\o  additional  sijilional 
membranes  e.ui  be  elev:ited  and  angled  toward  each  oth- 
er isolating  llie  pseiidolaixes  ami  functioning  a,s  a  txpical 
\vaste  caiKil.  :is  Kellogg  (1915)  descrilied  for  different 
species  of  bivalves. 

The  siphonal  meiiibr:ines  iiuiv  be  raised  lo  IreeK  ad- 
mit the  iuciirreiil  sli"e;im.  or  iiiav  be  drawn  {k)wnward 
to  direct  the  stream  low.ird  the  inantli'  edges,  and  away 
from  the  gills,  such  as  when  much  sediment  is  present. 
In  this  last  configuration,  a  ri'lativeK'  large  amount  of 
sediment  would  be  dejiosited  ventralK  on  the  mantle, 
and  i|iiicklv  lr:iiispiirled  posteiiorK. 

In  T  icnlricosa.  the  siphonal  membranes  control  the 
water  current  that  passes  through  the  mantle  caxitx; 
while  the  additional  folds  can  be  elevated  and  angled 
toward  each  other  until  tliev  meet  in  the  summit  of  and 
enclosing  the  inciirrent  siphonal  opening.  This  is  proi)- 
ablv  an  adapl:itioii  to  large  amounts  of  snsjiended  sedi- 
ment in  the  water  bv  directing  pailicles  awa\  from  the 
ctenidia  and  aiding  in  tluir  rcmoxal. 

.As  seen  in  some  N'eneridae  (.\nsell,  1961),  the  tree 
edge,s  of  the  m:mlle  li.ive  lour  lolds.  .According  to  Yonge 
(1957)  and  .\nsell  (1961).  the  presence  of  four  folds  is 
accomplished  In  the  duplication  of  the  middle  mantle 
fold. 

.\  large  n ber  ol  brancheil  tentacles  in  the  margin 

of  die  iiKinlle  was  described  tor  T.  cra.ssatdoidcs  (Kel- 
logg, 1915)  and  T  niactroidcs.  In  T  icnlricosa  there  are 
;ilso  branched  tent:icles  ])rolecting  the  organs  ol  the 
mantle  c:ivitv  Iroiii  inli-usion  of  large  particles  ol  sand 
and  possible  injiuv.  Manv  specimens  showed  torn  or  re- 
jxiired  oiilei  (leiiiibi'.mchs  as  apjiarent  conse(|iiences  of 
such  injnn,  as  (aien'in  and  Narchi  (2000)  also  siiowed 
for  I'rololhnca  ( Lcnkoniai  pcclorina  (Lamarck,  1818). 

The  surface  of  the  mantle  in  T.  vcntiico.ta  has  \-en- 
trallv  direcled  cle:iiising  cilian  ciuTent.s.  similar  to  that 
observed  bv  .\iisell  lOOL  in  soni<>  Wneroidea  fnjm 
Kngland  and  in  T  inaclroidcs.  In  T  icnlricosa  aiul  T 
niaclroidcs.  minor  cilian  cleansing  currents  pass  from 
the  free  margin  of  the  mantle  in  the  posterior  region, 
passing  radialK  inward  to  join  the  main  rejection  tract. 

.Ansell  (1961)  described  some  \ariation  in  outer  de- 
mibranch  confignration,  betxveen  species,  and  e\-en  be- 
txxeen  specimens  of  the  same  species  due  to  the  strength 


Pa-e  22 


THE  NAUTILUS,  \ol.  116,  No.  1 


sa6 


af 

Figure  16.  Tivda  ventrirosti.  Interior  of  stonuicii.  seen  From  rigiit  side  alter  opening  bv  ineision  in  tiie  right  wall:  rif.  anterior 
lokl:  lis,  beaded  swellings;  chl.  digesti\e  diverticnla;  (III.  dorsal  Jiood;  c,  esophagns:  gs,  gastie  shield:  ig,  intestinal  grooxe:  Ir.  left 
caecum;  /;;,  left  pouch;  ml,  minor  t\phlosole:  n:  right  caecum;  nii.  mouth  ring;  sr/,.  principal  sorting  are;i  ot  the  d(>rs;il  hood;  sii,„ 
sorting  area  of  the  lel't  pouch;  «;-,  sorting  area  below  esophageal  orifice;  .v.v,  style  sac;  ty.  t\plilosole. 


of  tlip  current  at  the  free  edge  earning  particles  tow  aids 
the  palps.  .According  to  Ansel!  (1961),  Vcocnz/j/s  aitrca 
(Gmelin,  1791  .  \'(>unij)is  rhoinhoidcs  (Pennant.  1777). 
Mijsia  innlfila  Peimint.  1777)  and  members  of  Vt';i(/\ 
Linne,  175S  aiit;  Di'^iuia  Scopoli,  1777,  present  an  in- 
cipient current  directed  toward  the  month  at  tlie  free 
edge  of  die  oiiii  i  de.rl'DJiincli. 

The  pattern  and  general  paths  ol  the  etenidiacnrrents 
of  T.  ventricosa  are  i:;  agreement  witli  the  results  ol 
Ridewood  (190.3)  and  Atkins  (19.37a.  b)  on   the   mor- 


[)hol(ig\  of  the  ctenidia  :uid  their  cihation  in  some  \'e- 
neroidea.  Tlie  ctenidia  in  T.  ventricosa  belong  to  T\pe 
V.{\)  of  .'\tkins  (1937b\  with  one  groove  along  the  free 
ventral  margin  of  the  iniu-i'  deniibranch.  as  \;ui'hi 
(1972)  and  Xarchi  and  Ciibrii'lH  ( 19S()>  described  for  tlie 
venerids  Aiioniahx'iinlid  hnisiliana  iCmelin,  1791)  and 
Chionc  suhroslrdid  (Lamarck,  1818).  respectively.  In  T. 
Iliad roidcs  the  eteiiitli;il  eiliation  is  of  T\pe  C(2)  of  At- 
kins i  1937b). 

The  hibi;il  palps  of  T  i ml ricosn  sliiiw   the  s;mie  b;isic 


W;  Xairlii  and  F.  di  Dario.  2002 


Paw  23 


features  of  ntlier  species  oIN'eneroidea  studied  In  Tliiele 
(1886),  and  are  also  similar  in  stnieture  and  niuseular 
acti\it\  to  tliose  of  T  iiiiictroidf.s.  The  \(-ntr,il  tips  of  the 
most  anterior  hlann'iits  of  tlie  inner  demiliraneli  are  in- 
serted into  and  fused  to  a  distal  oral  groo\c:  the  asso- 
ciation of  ctenidia  and  laliial  palps  belong  to  ( ^alegon  II 
of  Stasek  (1963). 

The  configuration  of  the  alimentan  canal  of  7'  icn- 
tricosn  is  similar  to  that  of  T  niiutrnidcs.  differing 
uiainK  In  the  nioic  coii\oluted  midgut,  with  5  closeK' 
packed  coils  in  the  first,  anil  with  oiiK  one  loose  coil  in 
the  latter. 

\\'here\er  known,  the  aiiatomx  of  the  stomach  is  gen- 
eralK'  similar  throughout  the  X'eneridae.  The  left  and  the 
right  caeca  of  T.  ccnthcosa  receive  4  and  6  ducts  from 
the  cligesti\e  di\erticula.  respecti\elv.  Six  and  .5  ducts 
were  respectixek'  recordetl  for  T  niactnudcs  In  7'  vni- 
tricosa  another  group  of  fi\e  tlucts  from  the  digestive 
diverticula  enters  the  li'ft  poucli  without  being  associ- 
ated with  the  major  t^plilosole  or  the  intestinal  gnxne, 
as  described  h\  Furchou  1 19fS())  for  C.ajninum  iniiiiniiiin 
(Montagu,  fS47)  and  7'  iiiiictniiilcs. 

The  stomach  of  7'  iciitiifasd  has  the  structmc  of  a 
tvpical  suspension-feeding  cnlamellibranch.  It  is  similar 
to  those  of  the  vfuerids  Olossus  liuiiKniWi  (I.-inne,  1758), 
G.  iniiiiviiiin.  \Cnii\  ciisiiKi  Linne.  1758.  and  7"  iiKKiro- 
idi'.s,  as  described  In  Owen  (1953).  Purchon  (1960).  .\u- 
sell  (1961),  and  Narchi  (1972),  respectiveK.  Within  the 
stomach,  food  particles  are  kept  in  motion  In  thi'  com- 
bined action  of  the  rotating  cnstalline  stvie  and  the  cil- 
iated walls,  and  are  subjected  to  sorting  maiuK  in  the 
posterior  sorting  area,  as  in  T.  luactroidcs.  The  stomacli 
of  T.  ventricosa  luav  be  capable  of  handling  luanv  par- 
tick'S  at  one  time  diii'  lo  lis  complexity. 

.AC'KXOWLKDC^MUXTS 

To  Conselho  Xacional  de  I.5esen\c)l\imento  (,'icutifico  e 
Tecnologico  (CNPf|).  Bra/il.  for  the  Support  (irant  nniii- 
ber  300490. 


LITER.ATUHK  (TFKD 

Alilidtt.  1-i,  T  U)74  .\iiiriR'aii  .Seashells.  2ikI,  Kdilion  \an  \i)s- 
traiid  l^einliold.  New  York.  663  pp. 

Ansell,  -\.  D.  1961.  Tlie  tiuictional  rnoipliologx  ol  Bnlisli  spe- 
cies of  \'eneracea  (Eulaniellihranchia).  Joiini.il  ol  llir  M.i- 
rine  Binlotjical  .\ssociatiiiTi  (if  the  United  Kiii^diiiii  )li2  : 
4Sy-.517 

Arinstrcina;,  L.  H.  i9(i.5.  liurniwiiin  liiiiitatidiis  in  I'llcrspnda. 
The  Wliiier  7:  19,5-2I)(1, 

Atkins.  D.  1937a.  On  the  cilian  inechanisnis  and  interrelation- 
ships of  laniellii)ranchs.  Part  II:  Sortint;  devices  on  the 
gills.  QuarterK- Journal  ol  Mitroscopical  Science  N..S.  79: 

Atkins.  D.  1937li,  On  the  eilian  mechanisms  and  interrelation- 
ships of  Lamelliliranch.  Part  III:  T\pes  of  fnnellihraneii 
gills  and  their  food  currents.  Quarterl\  Jdurnal  ol  Micro- 
scopical Science  N.S.  79:  37.5-421. 

Broggi,  J.  1970.  Sobre  el  hallazgo  de  ciuco  nue\  as  especies  para 


el  Qnerandino  Unignavo.  Comunicaciones  de  hi  .Soeieckid 
.\I:ilaeoloiiiea  del  Uruguay  2iIS»:  427—131 

Hneku[i.  I.,  19.57.  C'atalogo  dos  moliiscos  do  .\Iusen  Piiim:m- 
(k'lise  de  ( liencias  .Naturais.  Ilieringia  1:  1—10. 

Carcelles,  \  K  1944.  (.'atalogo  de  los  moluscos  marinos  de 
Puerto  One(iuen.  Re\ista  del  Museo  de  Pa  Plata,  \ne\a 
Seiie.  Zoolot;ia  3:  2.53~.509. 

Carcelles.  A.  H.  and  S.  1.  Williamson.  1951.  Catalogo  de  los 
moluscos  marinos  de  hi  Prorincia  Ma<;,illaiiica.  Ke\istadel 
Instituto  Xacional  de  Investigacion  de  las  (  ienc  i:is  Natnr- 
ales2l5):  22.5~.3S3. 

Coan.  K.  \'.  I'  Nalenfich  Scott  and  F.  R.  Bernard.  2000.  Bi\aKe 
seashells  ol  western  North  .America.  Marine  l)i\alve  mol- 
liisks  From  .\rctic  .\laska  to  Baja  California.  Santa  Barbara 
Museum  of  Xatunil  Histon-  Mono<;rapli  2,  Sant:i  Barbara, 
\iii  +  764  pp. 

Golterje,  C.  .\.  1950.  CJoutnhui^ao  a  zoosjeografia  ila  makico- 
fauna  do  litoral  do  estado  do  Parana,  .-^ninivos  do  Museu 
Paranaense  8:  221-2S2. 

Gneron.  C.  O.  C.  and  W.  Narchi.  2000.  .\natona:i  luncional  de 
Prototluicii  (Lctikttnui)  pectunna  (Lamarck.  ISIS)  (Bi\al- 
\na:  Wneridae).  Re\ista  Brasileira  de  Zoologia  17:  1007- 
I  ( i:  >9 

Kello'.;^.  I  I,,  1915.  ( !ilian,  mechanisms  ol  lamellihranchs  with 
desenptiiiii  (j|  anatonn.  journal  ol  .Moi]iIiolog\  26:  62.5- 
701. 

Lauije  de  .Morretes.  F.  1949.  Ensiiio  tie  cat:ilogo  dos  moluscos 
do  Brasil.  ,-\njiii\os  do  .Museu  Paranaense  7(1):  .5-216. 

Narchi.  W.  1972.  Comparati\e  stndx"  of  the  hmctioiuil  mor- 
pliolot^  ol  AiuiiiudiHiirdiii  hrasiliiinii  ((anelin.  1791  :ind 
Tivcld  iiinciroidi's  (Born.  177S!  (BivaKia.  \cneridae  Bul- 
letin ol  Marine  Science  22:  643-670, 

\:irc  111.  W,  and  M.  .\.  CJahiielli.  19S0.  Sobre  aiiatomia  lunc'ional 
de  CliioiK  stdirostratii  (Lamarck,  ISIS).  Re\ista  Xordes- 
tina  de  Biologia  3:  25-46. 

Narchi.  W.  and  S.  G.  B.  C.  Lopes.  I99S.  On  the  meiirreiil 
siplional  membrane  oiTivcla  vcniricoso  (C^raw  IS3St  (Bi- 
vaKia: \'eneridae).  Jn:  Bielen  R  and  P.  M.  .Mikkelsen 
(eds.)  .-\hstracts  of  the  World  (a)ngress  of  .\lalacok)gv. 
Washington.  D.C..  PJOS.  Unitas  -Malacolos;ica  and  .\mer- 
icaii  Makuologieal  .So(ietx.  \\'asliini;ton.  p.  2.37. 

Owen.  G.  19.5.3.  On  the  hiologv  ol  Glitssiis  humiinus  i  L.  i  i/.v- 
ociirdki  cor  Lam.'.  |oiinial  ol  the  Marine  Biolosjical  .Vs- 
.sociation  ol  the  United  Kingdom  .32:  S.5-106. 

Purchon.  R.  I).  19(iO.  The  stomach  in  the  Eiilamellihr.iniiiia: 
stoiiLieli  l\|ies  l\  .iiid  \.  Proc(>e(lings  ol  the  /.oolot;ical 
Sotielv  of  London  1.35:.!):  431-4.S9. 

Ridewixid.  W  C  190,5,  On  the  structure  ol  die  >;ills  ol  the 
Lamelliliraiicliia.  Philosophical  Transactions  ol  the  Ro\al 
Societv  of  Loiukin  Ser.  B.  195:  147-2S4. 

Bios.  I'',  (.'.  1970.  (aiastal  Brazilian  Seashells.  I'uiidavao  Cidade 
do  Kid  Gr:iiide,  Fundayao  I'nivcrsid.ule  do  Rio  (iraiide. 
Rio  Cr.iiide.  2.55  pp.  60  pis. 

Rios.  L.  C.  1975.  Bra/ili:in  Mariiii'  Mollnsks  Iconoiiiaphv.  I'iiii- 
dayao  Universidade  do  Rio  (inmde.  Rio  (a'aiide.  331  pp.. 
91  pis. 

Rios.  F.  C.  19S5.  .Seashells  of  Bni/if  Isl,  edition.  l-iiiida(,'ao 
Cidade  do  Rio  (iraiide.  F'uiida(,ao  Universidade  do  \\w 
(Jrande,  Miiseu  Oceaiiografico.  Rio  Grande.  32S  pp.,  102 

Rios,  L.  C.  1994.  Se:isliells  ol  Brazil,  2iid,  edition,  l-undavao 
Cidatk'  do  Rio  Grande,  Instituto  Ac()ua  R.J..  Museu 
Oceaiiografico  Prof  E,('.  Rios.  Univ(>rsidade  do  Rio 
Grande,  Rio  Grande,  .36S  pp.,  I  13  pis. 


Paee  24 


THE  NAUTILUS,  \ol.  llfi.  NO.  1 


StiLsek.  C,  H.  1963.  S\niopsis  and  cliscnssion  of  the  association 

of  ctenidia  and  labial  pal|)s  in  tlic  l)i\aKcd  Mollnsca.  The 

\eliger  6:  91-97. 
Thiele,  J.  1S.S6.  Die  Mundlappen  der  Laiiullihi.uichiaten.Zeit- 

sclirif't  fiir  wissenschal'tliclie  Zoologie  44:  239-272,  2  pLs. 
WVviiiouth,  F.  W.  1920.  The  edible  elanis,  nuissels  and  scallops 

of  Calilornia.   Hnlletin  ol   the  (California  State   Fish  and 

Game  CConi  mission  4:  3-72,  19  pis 
Wevmouth,  F.  VV.  1923.  The  lile-histon  and  u,ro\\lli  ol  the  IMs- 


mo  clam  {Tirclti  stitllonitn  Mawe).  Bnlletin  of  the  Cali- 
fornia State  Fish  and  dami'  Connnission  7:  5-120. 

Von<j;e.  (;.  .\1.  f948.  Formation  of  siphons  in  I>amellibraneliia. 
"  Nature  161:  198-199. 

Yonsie,  (;.  M.  1957.  Mantle  fusion  in  the  1  ,ainellibranehia.  f'ub- 
blica/.ioni  de  la  Sta/ioue  '/ooiot;ica  di  Napoli  29:  151-171. 

Yonge,  C  M.  1982.  Mantle  inar<;ius  with  a  re\'ision  of  siphonal 
t\pes  in  the  Bi\aKia.  |onru:il  ol  Miilluscaii  Studies  48: 
102-103. 


Notice 


THE  2002  R.  T.  .\BBOTT  VISITING  CURATOHSlilP 


Tlic  |-5ail('\ -Matthews  Shell  .Vluseuin  is  pleased  to  invite  applications  lor  the  2002  R.  T.  .\liliott  Nisitiin^  ( Jiratorsliip. 

The  Ciiratorship.  e.stablisliecl  oriwinallv  in  accordance  with  tlie  wislies  ol  the  late  Dr.  R.  Tucker  Abbott,  Foundint; 
Director  of  the  Shell  Museum,  is  awarded  annually  to  enable  mollusk  .s\steiiiatists  to  \isit  tlie  museum  for  a  period 
of  one  week.  .Abbott  Fellows  will  be  expected,  by  performing  collection-based  research,  to  assist  with  tlii'  curation  of 
porticjns  ol  the  Museum's  collection  and  to  provide  one  evening  talk  for  the  general  public.  The  Museiun  collection 
consists  of  Tuarine,  freshwater,  and  terrestrial  specimens.  A  large  percentage  of  oiu"  hoklings  have  been  catalogui'd 
tln'ougli  a  coniputerizeil  database  managi-ment  svstem.  .\  substantial  portion  of  the  time  will  be  available  for  research 
in  the  colk'ction,  but  field  work  in  southwest  Floritla  can  be  arranged.  The  R.  T.  Abbott  \isiting  (!iu-atorship  is 
accompanied  bv  a  stipend  of  $1,500. 

Interc\sted  malaeologists  are  invited  to  send  a  copv  ol  tluir  ciiiiieuhnn  vitae  together  with  a  letti'r  tletailiu'j;  their  areas 
of  taxonomic  e.xperti.se  and  resi.'arch  objectives,  and  to  pinvide  a  tentative  subjei-t  for  tluir  talk.  Si'ud  materials  to: 

Dr.  Jose  11.  Leal,  Director 

The  Bailev-Matthews  Shell  Museum 

P.O.  Box  ioSO 

Sanibel,  Fi,. 3.3957 

jleal@shelhmiseum.org 

.Applications  for  the  2002  X'isiting  Curatorship  should  be  sent  no  Liter  than  .Mav  .30.  2002.  The  aw:nd  will  be  aimouni'ed 
by  late  June.  Questions  about  the  \'isitin<j;  ( 'malorship  slmnld  be  sent  to  tlie  e-mail  address  above,  or  liv   piinne  at: 

(941)  39.5-22.33:  lax  (941)  .395-0706. 


THK  NAUTILUS  1  Ifii  ]  ):25-:3I.  2002 


Paw  25 


Use  of  HMDS  (hexamethvldisilazane)  to  dn-  omiiiic 
microstructures  in  etched  bixaK e  niollusk  and  barnacle  shells 


Beind  R.  Schone' 

Dt'purtiiieiit  of  Geosciences 
Universitrv"  of  Ariz.ona 
Tucson,  AZ  85721  USA 
l)enKl..schoene@excite.coiii 


Da\id  Bentley 

l5i(it('(liiioloin,-  Imaging  Facilit\ 
Uiii\cisih  ol'  Arizona 
Tucson,  A/  S5723  USA 


ABSTRACT 

The  organic  framework  of  molfnscan  and  iiarnacie  sjiclls  \ields 
clues  to  Ijiocalcification  processes.  Sliglit  demineraiization  of 
the  shells  reveals  the  fragile  meshwdrjs  of  insoluble  organic 
fillers  and  membranes,  which  tend  to  collapse,  wrinkle,  and 
shrink  when  air-dried  from  water  Gomparison  of  different  diT- 
ing  teclmiijnes  on  etched  bi\al\e  mollnsk  iChionc  flti(tifr(i'j,ti) 
and  baniacle  shells  (Chtlunudlus  sp.)  reveals  that  hexanietli\kl- 
isilazane  (HMDS)  produced  results  qualitatiscK  superior  to 
critical  point  dning  or  dning  from  eth\l  alcohol  or  water 
H.MDS  dries  stnictural  details  of  the  organic  iiicsliwork  e\ccl- 
lentK  and  facilitates  the  recognition  of  faint  growth  nicrements 
for  growth  pattern  analysis  (sclerochronolog\).  The  H.MDS 
method  is  cost-effecti\'e,  sa\es  time  and  can  lie  used  as  a  rou- 
tine substitute  for  drying  microstructures  in  slightK  etched 
molluscan  and  baniacle  shells. 


INTHODl  CflOX 

Organic  matter  plaws  an  es.sential  inle  in  the  formation 
of  molhi.scan  and  baniacle  shells.  It  has  been  realized 
that  the  organic  matri.x  proxitles  the  structural  frame- 
work ("template";  Clark,  1980)  for  hiomineralization  and 
influences  the  mineralogical  and  cA'stallographic  prop- 
erties (Mann,  1983:  Simkiss  and  \\'ilhur,  1989;  Cren- 
shaw, 1990  and  literature  therein;  W'atahe  et  al.,  1993). 
Demineraiization  of  skeletal  hard  parts  mixcils  the  un- 
derKing  three-dimensional  organic  microstrmtures, 
which  ma\'  vield  clues  to  hiocalcificatioii  processes. 

Organic  matter  is  also  a  major  constituent  of  some 
growth  increments  (e.g.,  Koike,  1986).  Slight  deminer- 
aiization of  cross-sectioned  molhiscan  and  InuiKicle 
shells  re\eals  a  three-dimensional  relief  of  insoluhli'  or- 
ganic components  and  dilferentialK  dissoKcd  cnstals  (as 
a  result  of  different  crxstal  sizes  and  orientation).  In 
sclerochronological  studies  (growth  ;iual\ses),  supei'fieial 
etching  is  commonly  used  to  aid  in  the  identification  and 
measurement  of  internal  growtli  ini'rements  in  niollns- 

'  Current  address:  Institute  and  Musiuni  lor  Ceologx  :ind  I'a- 
laeontologN,  |ohaiin  Wolfgang  (ioethc  L!in\ersity.  Senckeii- 
berganlage  32-34,  60054  Frankfurt  am  .Main,  (iermany. 


can  and  Inuiiacle  skeletons  (Khoads  and  Lutz,  1980; 
Scluine  et  al.,  in  pi'ess).  The  etching  time  varies  for  ilif- 
terent  species  and  depends  on,  hir  instance,  the  shell 
strnctinv,  mineralogv,  and  organic  content.  .Although 
growth  patterns  in  mollnsk  shells  are  the  focus  of  im- 
ineious  studies.  ouK  lew  papers  deal  with  the  growth 
patterns  of  harnack's.  Unliki'  most  crustaceans,  lianiacies 
do  not  replace  their  hardparts.  Both  mollnsks  (e.g..  Dav- 
enport. 1938;  Fannelhi  and  .MacCJlintock.  1968)  and  bar- 
nacles (e.g.,  Bourget,  1980)  grow  bv  periodic  accretion 
of  skeletal  material  producing  circadian  growth  incre- 
ments (see  sketches  in  Figure  1  and  2:  direction  of 
growth).  In  baniacles,  the  growth  lavers  are  best  \iewed 
in  the  she;itli  Liscr  (I'"igure  I),  and  in  most  bi\alM-  mol- 
lusks  in  the  outer  shell  laxcr  (Figure  2i. 

Like  most  solt  tissues,  the  shell  organic  framework, 
including  the  orgauii--rich  growth  increments  obseiM'd 
in  cross-sections,  is  jirone  to  collapse,  shrinkage,  and 
wrinkling  when  ;iir-dried  le.g.,  .Anderson,  1951;  Nation, 
1983;  Clark,  1980,  1999).  Hre\cnting  these  unwanted  ef- 
fects requires  special  cluMiiical  treatment,  which  (k>h\- 
drates  and  hardens  the  fragile  organic  structures.  Several 
techniques  are  used  to  di\  biological  solt  tissues,  .al- 
though extremeK  time-consuming  and  quite  dangerous 
(highl\-  pressurized  chamber),  critical  point  drviug 
(CPD)  is  b\  far  the  most  common  method  (Anderson, 
1951:  using  liquid  COj,  e.g.,  Clark.  1980  or  Freon  13  as 
a  transitional  fluid,  e.g..  Koik(\  1986).  On  average,  pre- 
paring one  sample  In  WD  re(juires  full  attention  o\er 
1.5  hours.  The  basic  (.'PD  e(juipment  costs  se\-eral  thou- 
sand dollars,  (iood  results  were  also  achiexcd  with  the 
sublimation  deli\di;uit  Peldri  II  ( K(>n!ied\' et  al..  1989). 
lIowcM'r,  prepar;itiou  lollowiug  this  technique  takes 
more  than  t^\■ice  the  time  as  (;PD,  and  Peldri  II  is  no 
longer  axaihible  because  of  environmental  hiizards.  Flu- 
ids with  low  surface  tension  (ac-etone  or  propxlene  oxide, 
Bovde  and  Wood.  1969)  sometimes  produce  reasonable, 
artifact-free  results  for  biological  soft  tissu(>s.  Some 
workers  prefer  the  extremeK  hazardous  osmium  tetrox- 
ide  technique  iOnattlebaum  and  Carner,  19801. 

.\  reliable  ;uid  sim]ile  dning  techni(|ue.  which  pro- 
duces   results    (|u;ilit;iti\el\    eompai:ible   or   superior   to 


Pac{e  2tt 


THE  \AUTILUS.  \<)1.  116.  Xo.  1 


2  mm 


2  mm     / 

\ 

/ 

cross-section 

'^ 

parietes 


Figures  3-5 


1  cm 


Fieures  14-16 


''^ 


inner  shell 
surface 


\ 


Figures  6-8 


commissure  line  (outer  rim) 


Figures  9-13 


cross-section 


B.  R.  ScIkhk- 


D.  Bcntlfx.  2002 


Paffl" 


CPD.  was  introduced  In  Nation  (19S3).  After  tlelndra- 
tion  in  a  series  ot  graded  ethanol  solutions,  tlie  samples 
are  innnersed  in  hexamethvl-1,  1.  1.  3,  3.  3  disilazane 
I hexanieth\klisikvzane.  HMDS),  an  organic  reagent  with 
tlie  clieniical  formula  [(CH5),Si],NH,  for  approxiniateK' 
fi\e  to  30  minutes,  and  allowed  to  air-dn'  at  room  tem- 
perature. The  low-cost  HMDS  dning  technifjue  does 
not  require  full  attention  during  processing.  It  was  suc- 
cessf ulK  applied  to  dn-  \arious  kinds  of" soft  parts  in  dif- 
ferent organism  groups,  e.g.,  in  insects  (Nation.  1983; 
Rumph  and  Turner.  199S).  soft  tissues  of  niollusks  (.Leal 
and  Simone,  1998),  \ertebrates  (Heegaard  et  al..  1986; 
\\'er\ha  et  al..  1990),  microorganisms  (Dekker  et  A.. 
1991;  Hochherg  and  Lit\aitis.  2000)  or  pollen  (Chissoe 
et  al.,  1994).  .\lthough  ;ipplied  to  deniineralized  human 
teeth  (Perdigao  et  al.,  1995;  CaniJlio  et  al..  1996).  to 
our  knowledge  HMDS  h;is  not  been  used  as  a  transi- 
tional soKent  for  dning  molluscan  and  baniacle  shell 
microstnictures. 

This  stniK'  e\aluates  the  usefulness  of  ll.MDS  for  lin- 
ing microstnictures  in  etched  barnacle  and  bi\ al\e  mol- 
lusk  shells  using  scanning  electron  microscopy  (SEM). 
Special  emphasis  is  gi\en  to  the  use  of  HMDS  in  scler- 
ochronological  studies.  We  compare  the  HMDS  tech- 
nique to  1)  the  CPD  method.  2)  ;iir-dning  immediately 
after  etching  (.\IR),  and  3)  a  combination  of  dehydration 
in  a  series  of  graded  alcohol  baths  and  ;iir-di"\ing  i.\LC). 

MATERI.\LS  AND  METHODS 

Material  and  general  preparation:  Durint;  field 
trips  in  1999  and  2000.  we  collected  biirnacles  and  bi- 
\iJ\'e  mollusks  in  the  intertidal  zone  of  the  northern  Gulf 
of  California,  Me>dco.  For  this  stud\,  we  used  one  bar- 
nacle specimen  (Chthainahis  sp.)  collected  ali\e  at  Isla 
Sacatosa  in  December  2000  (N31  29.60,  \M14  50.85; 
specimen  no.  ST11-A18),  three  Cltionr  fiiiclifrana  (Sow- 
erbv,  1853)  specimens  collected  ali\e  at  North  Orca  in 
November  1999  (N31  32.60,  WH4  52.78;  specimen  no. 
NO3-A105,  NO3-A106,  NO3-A108),  and  three  C.  fluc- 
tifms^a  specimens  from  Isla  Sacatosa  (N3I  29.60.  \\  114 
50..8.5;  specimen  no.  ST12-D1.  .ST12-D2.  ST12-D3;  De- 
cember 2000).  All  specimens  are  housed  at  the  Depart- 
ment of  Geosciences,  The  Uni\ersit)  of  .Arizona.  Clollec- 
tion  Flessa,  Subcollection  Schone. 

.After  remo\al  of  the  soft  parts,  all  specimens  were 
cleaned  in  Chlorox  (5.25'7r  soilium  Inpochlorite)  to  dis- 
sol\e  remaining  superficial  organic  matter,  rinsed  \\ith 
water  and  dehvdrated  with  ethyl  alcohol  (EtOH).  Coat- 
ing with  ]-B  KWIK  Wekr^'  pro\ides  a  protecting  co\er 
for  the  shells  during  cutting.  The  fragile  barnacle  re- 


(juireil  I'mbcdding  in  |-B"  Epow.  |-B°  Epow  is  not  ad- 
vcrseK  affected  b\  immersion  in  EtOH  or  H.MDS. 

Cross-sections  of  Ijarnaclc  and  lji>al\e  inolliisk 
shells:  We  obtained  three  cross-.sectioned  slabs  (thick- 
ness 0.5  uun)  from  the  barnacle  shell  (ST11-A18;  Figure 
1)  and  from  the  left  \aKe  of  each  of  the  three  C.  fluc- 
nfmoa  shells  no.  ST12-D1,  -D2.  and  -D3  (Figure  2)  us- 
ing a  low  speed  (Buehler'^'  Isomet')  saw.  Use  of  an  ul- 
tratliin  saw  blade  (0.4  mm  thickness)  assured  that  the 
effects  of  different  preparation  techniques  (see  below) 
on  the  organic  microstnictures  could  be  studied  on  id- 
most  identic;il  portions  of  the  shell.  In  this  stud\  we 
focus  on  the  composite  prismatic  shell  la\cr  of  C.  flttc- 
tifraga.  because  the\  are  of  primarv'  concern  in  sclero- 
chronological  studies.  In  the  cross-,sectioned  shells  of  C 
fnctifni'^a  growth  increments  are  clearK'  de\eloped  and 
easv  to  measiue.  Moreo\er,  etching  and  dning  results 
ma\'  \an'  with  respect  to  different  hpes  of  shell  inicro- 
stiTicture.  Focusing  on  one  tvpe  of  shell  la\er  allows 
comparisons  of  the  effecti\  eness  of  different  dr\ing 
techniques  on  similar  shell  microstnictures.  The  cross- 
sections  were  mounted  on  petrographic  slides,  ground 
on  glass  plates  (600  and  1000  grit  -AbO,,  powder),  and 
polished  on  laps  (9,  6,  and  0.3  |jLm  .Al^O,  powder).  Ul- 
trasonic cleaning  bet\\een  e;ich  polishing  step  assured 
that  no  grinding  powiler  was  left  on  tlie  surface. 

Inner  and  outer  sliell  suriaces  of  the  bivalve:  Shell 
fragments  of  three  C.  Jiiictifraga  specimens  (NO3-A105. 
-.A106.  and  -.A  108)  were  mounted  on  a  petrographic  slide 
to  expose  the  growing  surface  at  the  inner  shell  edge. 
Two  shell  fragments  of  specimen  ST12-D3  were  mount- 
ed on  petrographic  slides  with  die  outer  surface  exposed. 

Chemical  treatment:  etching  and  dicing:  All  sam- 
ples were  then  etched  in  a  0.25  inol  EDT .A  solution  (eth- 
\lene  diamine  tetraacetate.  pH  7.95.  buffered  with 
NaOH)  and  carefulK  rinsed  in  de-ionized  water  The 
shells  w-ere  etclied  for  \aning  amounts  of  time  (Table 
1 )  to  obtiiin  different  degrees  of  demineralization:  shglit 
supei-ficial  etching  and  decalcification  of  the  upper  1  mm 
of  shell  niat(Mial.  In  previous  studies  we  canied  out  a 
series  of  tests  and  anakyed  the  effects  of  an  array  ot 
etching  times  and  EDTA  concentrations  on  shells  of  C". 
j{uctifm<i(i  and  Chthainahis  sp.  (Schone  et  al.,  2002.  and 
unpublished  data).  For  the  presentation  of  microstnic- 
tures in  C.  fiuciifraga  and  Chtliaiiuihis  sp.  the  approach 
used  herein  pnned  to  be  most  appropriate.  We  com- 
pared the  following  lour  dr\ing  techniques  (Table  1);  air- 
dning  from  water  (AIRi.  dning  from  EtOH  (.ALC).  dn- 
in<r  from  H.MDS  and  CPD.  For  ALC.  HMDS,  and  CPD 


Figures  1.  2.  Positions  in  the  shelLs.  wliere  tlie  samples  (Figures  3-16 1  were  taken,  d.^s;  =  direction  ot  growth.  cUrecfion  ,n  winch 
sulisequent  crrowth  increments  were  added.  1.  Cross-section  through  the  p;iri.-tes  (shell  plates)  of  tiie  lianuicle  Chthamalu.s  sp.  The 
sheath  laver  (distal  portion  of  the  parietes)  exhibits  faint  dailv  growtli  stnictiires  T-igiires  .3-.5).  2.  Chiomj{m-t  if rn<ia  sheW.  Samples 
for  Fienres  6-S  were  t;iken  from  tlie  grouiiii;  edge  of  tlie  inner  shell  surface,  samples  for  Figures  14-l(i  from  tlie  outer  sliell 

lecome  appan-nt  in  the  outer  shell  laser  (Figures  9-13). 


lor  rignres 

surface.  In  radiiil  cross-sections,  growth  p.itteins 


Pa.'e  2S 


THF  NAUTILUS.  \()1.  116.  NO 


B.  R.  Stlioiie  and  D.  Bentlex.  2002 


Page  29 


the  samples  were  rinsed  in  a  sfrit's  of  graded  EtOH 
(40%.  BC^f,  80f7r,  5  min  each;  hvo  times  lOO'-L  30  iiiin 
eacli).  The  HMDS  technique  inchides  iiniiicrsion  ol  the 
dt'h\ih'ated  samples  in  97'^?-  hexametln  Idisilazane 
lAldrich'^'  chemical;  two  times  for  30  min  each!  and 
subsecjuent  air-dning.  CPD  was  performetl  in  ;i  Polaron 
CPD  E32()0  apparatus.  The  delndrated  samples  were 
placed  in  the  CPD  apparatus  with  alcohol,  the  chamber 
cooled  to  10°C,  and  CO^  allowed  to  enter  the  chamber 
Free  alcohol  was  flushed  from  the  chamber  until  dn'  ice 
was  seen  exiting.  The  sample  remained  in  CO,  for  45 
minutes  flushing  the  chamber  e\"en'  15  minutes.  At  the 
end  of  the  exchange,  the  chamber  was  sealed  and  the 
temperature  raised  above  35°C  (critical  temperature  for 
CO.  =  32°C).  CO^  \-apor  was  then  released. 

Coaling  for  SEM  and  photographic  documentation: 

.Vlimiinnm  stubs  were  glni'd  to  the  pctrogr;iphic 
shdes.  The  samples  were  sputter  coated  immediateK 
with  a  30  nm  gold  la\er  in  a  Bio  Rad  Polaron  Di\ision 
SEM  coating  svstem.  The  samples  were  stndietl  with  a 
Cambridge  Instruments  Stereosciin  120  SEM  at  accel- 
eration x'oltages  of  15  and  25  ke\'  Black  and  white  pho- 
tographs were  taken  on  Polaroid  Positi\"e/Negati\e 
4X5"  T\pe  55  Instant  Sheet  Film  toi-  ilocumentation. 

RESULTS 

Barnacle  shell  microstructure:  Under  750X  mag- 
nihcation.  the  15-min-etched  cross-section  of  the  sheath 
[lortion  of  the  barnacle  parietes  (Figure  I)  sliows  partly 
dissohcd  calcite  cnstals  in  a  crisp  three-diniensio]ial 
mesh  of  organic  fibers  when  dried  from  li.MDS  (Figure 
3).  CPD  results  in  a  less  well-dried  organic  framework 
(Figure  4).  Apparently,  some  delicate  organic  fibers  col- 
lapsed resulting  in  a  slightK'  bhurcd  image.  Even  worse 
is  the  drying  in  ALC  samples,  l^resniiuibly.  as  a  result  of 
a  complete  collapse  of  the  organic  iiiatiix,  single  ciystals 
can  bareh'  be  discerned  (Figure  5).  Some  growth  layers 
disappear  parth  or  entirely  when  immediateK  air-dried 
after  etchiuE;  (Fiiiure  5). 


Bi^al^e  shell  microstructure: 

Inner  shell  sire.vce  near  nii  <  ommfsm  ki  :      I! \il )S 

treatment  of  the  slightK  etclieil  growing  surface  at  the 
iimer  shell  surface  (Figure  2!  reveals  a  thin  organic 
mesliwork  residue  in  between  ;md  on  top  of  the  polyg- 
onal pattern  of  the  composite  prismatic  stnictnre  (Fig- 
ure 6).  CPD-treated  shells  are  not  depicted,  because  the 
results  are  companible  to  the  HMDS  d(>siccation.  .\L(J 
produces  a  soiuewhat  collajised  oi'ganic  framework  (Fig- 
ure 7).  The  rod-like  surface  is  compIcti'K  coxcred  in 
.\IR  samples  (Figure  S). 

CROSS-SEfTlON,  Ol  TER  COMPOSIl  E  PRISNLVIIC  SMELL 
UWER:  .\  cross-section  etched  for  15  min  and  dried 
fn)m  H.MDS  clearK  shows  the  composite  prismatic 
structure  of  the  outer  shell  layer  iFigures  2.  9).  Organic 
membranes  of  daih'  and  even  subdaiK'  growth  layers  are 
intersected  b\  peipendicular  running  fibrous  mem- 
branes (Figure  9;  for  exiierimental  studies  on  the  timing 
of  growth  pattern  foiination  in  ('  flii(iifra<ia  see  Scluine 
et  al..  2002).  CDP  dries  the  organic  mesliwork  less  well 
(Figure  10  i. 

.\t  lower  magnification  (375Xi  the  differences  be- 
txveen  the  HMDS  (Figure  11  >  ;md  C:PD  i  Figure  12) 
techniques  are  less  olnions.  .\  reli;ible  comitini;  ;mcl 
measurement  of  narrow  dailx  growth  increments  laid 
down  during  liot  sunmier  conditions  can  be  conducted 
on  HMDS  ;ind  CPD  s;implcs.  but  b;ircl\  accomplished 
on  die  sample  treated  with  the  .M.C  method  (Figure 
13!.  In  the  latter  case  the  collapsetl  organic  matter  cov- 
ers most  parts  of  the  faint  growth  patterns  ;ind  does  not 
allow  for  detailed  growth  pattern  ;inalysis. 

Outer  shell  surf.voe:  Outer  shell  surfaces  exposed 
for  t\velve  hours  to  the  0.25  mol  buffered  EDT.\  solu- 
tion reveal  a  well-dried,  dense,  nnilti-layered  organic 
network  when  dried  from  H.MDS  i  Fignr(>  14).  Growth 
lines  cannot  be  seen  iin\  more,  .\pplying  the  .MR  meth- 
od, the  organic  fnunework  collapses  and  covers  the 
shell's  microstructures  as  a  mass  without  clear  structures 
(Figure  15).  Durintj;  the  vacuum  in  the  sputter  coater 


Figures  .3-16.  SENi  iscanniixj;  electmii  iiiicroscopv  )  inuincs  iil  .tclieil  hdn\M\f  •ClillKnndlii-,  sp.)  (  Fis^ires  3-5)  and  bivalve  moliusks 
shells  (Cliit'iiv  fiuctifraoa)  (Figures  (i-t6).  Working  (list;iTice  in  iniu  iWDi.  atccleratioii  vnltasje  in  ke\'  l.W),  ptcliing  time  in  min 
(ET),  and  scale  are'provided  in  cacli  fit;iire.  .•}.  Best  diving  of  the  orijanic  mesliwork  (\>  and  single  cnstals  (cs)  is  achieved  when 
the  etclied  barnacle  shells  are  immersed  in  H.MDS  prior  to  air-diviim.  I'mminent  lid-ies  are  daih  iiiwvtli  lines  (dgi!.  4.  CPD  (critical 
point  dniiia)  dries  the  organic  network  paitlv.  hiil  the  sinjile  ciyst;ils  cannot  be  discerned  well  .5.  .Ml  structural  details  are  gone 
when  using^'ALC  (;iir-dning  from  cthvl  ;ikohi')b.  Some  faint  Growth  lines  iell  over  to  iidjacent  srrovvth  lines  (arrow).  6.  The  grovving 
edge  of  the  inner  suriac*  exliihits  crisp  poKnoiuil  cnstal  structures  in  the  H.MDS  treateil  specimen.  7.  CPD  dries  most  of  die  faint 
structures  as  well.  8.  ALC  results  in  a  blurred  image.  9.  Dailv  and  subdailv  (sgi)  sriovvth  increments,  organic  membranes  (onii  as 
will  as  peipendicular  ruimint;  orijanic  fibers  (of)  are  well  dried  in  H.MDS-treated  cross-sections  aU'Mmc fluctifras.(L  10.  but  less 
well  111  CPD  samples.  11.  Daih'  <;iovvth  lines  (diji)  laid  down  iliiriii-.;  hot  smuiiier  ;iie  best  viewed  in  HMDS  or  12.  CPD  prepared 
cniss-sections.  1,3.  ALC  does  not'allow  for  a  detailetl  analvsis.  because  organic  structures  colkipse.  14.  Outer  shell  snriaces  of  C/n«/K' 
fillet  if rw^a  reveal  a  dense,  three-dimensional  insoluble  cailaiiic  meshwork  when  immersed  in  HMDS  pnor  to  air-dning.  1.5.  .-MK 
(air-dnine;!  results  in  an  undifferentiated  mass  of  collapsed  oiaaiiic  stnictures.  16.  \\\mS  treated  samples  mav  shrink  and  wrinkle 
under'vacuum  during  the  coating  process  or  in  the  SEM  dies  =  non-etched  slu  II  siiri;ice.  don  =  demineralized  organic  network). 


p.  me  30 


THE  NAUTILUS,  \ol.  llfi.  \( 


Table  1.   0\enie\v  ol' s;imi|)1c  licatmcnt. 

l'R'ser\atioii  t 

ecliuiinie 

Species                   Spi'eliiirn  #            l-'tcliinjj;  time 

HMDS 

CPD 

ALC 

MB 

Chllwlamiis  fastis 

ST  11 -A  18 

13  mill 

Figure  3 

Chione  jiuc1ifr(i<!fi 

N03-A1().5 

3  mill 

Figure  6 

NO3-A108 

3  mill 

NO3-A106 

3  mill 

ST12-D1 

15  mill 

Figures  ^ 

STliTW 

12  li 

Fii'ures  1 

Fisrure  4 


9,  11  Figures  10,  12 

14,  K-i 


Figure  5 
Figure  7 
Fiiiure  13 


Figure  S 
Fisiiire  15 


and  the  SKM,  however,  the  organie  nehxork  .slieet  wrin- 
kles and  shrink,s  (Figure  16j. 

DISCUSSION 

Acl^anla^es  of  llio  HMDS  leclinitjue:  Iniinersion  in 
II.MUS  prior  to  air-(h"\iiig  th'ie.s  more  clearlv  the  true 
arrangement  of  the  insohihle  organic  microstmctures  in 
etelied  l)i\al\e  inollusk  and  liarnaek'  sheiks  than  ck)es  the 
eomnionK'  used  (,'PD  teehniijue.  CieneralK,  the  organic 
h-amework  of  slightK'  etched  cross-sections  or  etched 
surface  samples  is  well  presencd  in  its  original  three- 
dimensional  condition  when  dried  lioiii  HMDS.  This 
dning  (jnalit^•  allows  high  precision  anaKsis  of  the  struc- 
tural organic  framework  in  the  shells. 

CJrowth  patterns  in  IIMDS-treati'd,  slightK  deniiner- 
ali/.ed  shells  appear  innch  clearer  than  in  AIR  or  ALC 
l)rocessed  specimens.  In  the  latter  cases,  narrow  growth 
increments  are  eonipletel\-  covered  l)v  collapsed  organic 
material.  Under  the  hiirdeu  of  the  collapsed  organic 
incshwork,  fine  and  organie-rich  iiicrcinents  ma\'  fall 
ox'er  to  adjacent  growth  increments.  Tiicsi'  ad\erse  ef- 
fects prexent  sclerochronologists  from  counting  and 
measuring  the  growth  increments  correctK.  In  speci- 
mens dried  from  IIMJ^S,  e\'en  snh-daiK  grnwili  patti^ns 
can  be  ri-cogni/ed  with  ease. 

II.VIDS  dr\ing  does  not  re([uire  I'xpensivc  lab  e(|nip- 
mcnt  or  additional  skills  like  CTD.  Sample  preparation 
is  ea.sy,  takes  only  a  few  minules  and  does  not  re(|iiirt' 
constant  monitoring  during  the  immersion  or  the  dning 
process.  Moreoxcr,  the  sample  size  is  not  restricted  In 
the  small  vacuum  chamber  like  in  the  CPI)  apparatus. 
but  only  by  diffusion  constraints.  II.VIDS  dmng  is  less 
dangerous  than  CPD,  becau.se  it  (kies  not  re(|nire  high 
pressures  ilining  ]-)rocessiug.  which  ma\'  represent  some 
potential  vonrc(   of  hazard  to  the  operator. 

Disadvanlages  of  ilic  FiMDS  technique:  Ihick  or- 
ganic 1,1'. I  is  shrink  and  uriiikle.  These  artifacts  max  re- 
sult Iroi.i  the  :,nmersion  in  HMDS  itself  or  from  the 


high  xacuiiui    '..;,•: 
lem  is  also  v  i-m 
soft  tissues    f  '■    ,  ' 
ditional  fixatm 
ingand  HMDS    ; 
organic  mesliwork    i 
maliu  re|i()rtedK   ha 


•  caimg  and  SEM.  The  latter  prob- 

X.:  1.  r  CPD  and  Peldri  II  treated 

■  ■  '    .!..   llWfii.  It  is  not  clear  if  ad- 

^ .  iii   iriatri.x  prior  to  delivdrat- 

'     ■  III  staliilize  and  harden  the 

'!  ..  'I  liniiian  dentine  with  for- 

s    ;  '  ■    -I    I'lli'cts  on  the  stnicdiral 


arrangement  of  the  organic  fabric  (Canalho  et  ak,  1996). 
We  assume  that  longer  infiltration  and  dning  times  max' 
reduce  the  aforementioned,  unxvanted  effects. 

HMDS  is  a  hazardous  chemical.  Its  potential  effects 
on  humans  and  on  the  enxironment  are  barelx'  knoxxn 
(Material  Safety  Data  Sheet).  It  is  stronglx'  adxised  to 
use  the  chemical  onK  under  a  fume  hood  and  to  prexent 
inhahng  or  skin  contact. 

CONCLUSIONS 

HMDS  dning  is  a  xen-  useful,  loxx-cost,  easx-to-appK 
and  time  saxing  alternatixe  to  other  knoxxii  dning  tech- 
ui(jues  for  soft  tissues.  This  publication  demonstrates  the 
applicability  of  this  fast  dning  solxent  in  sclerochronol- 
og\'  and  for  the  studx'  of  organic  microstructm-es  in  some 
bivalxe  mollusk  and  barnacle  shells.  With  H.MDS.  xxe 
aehiexed  results  ([iialitatixelx  superior  to  the  xxidelx'  ap- 
plied CPD.  Further  studies  should  determine  if  the 
HMDS  method  can  be  used  as  a  routine  teclmicjue  in 
dning  organic  stnietures  of  etched  mollnsean  and  bar- 
nacle shells  and  if  the  HMDS  method  can  also  be  ap- 
plied to  etched  accretionan  hard  parts  of  other  marine 
or  terrestrial  organisms. 

,'U:kn{)\\lei:)C.ments 

This  studx  has  been  made  possible  bv  a  postdoctoral 
seholarshi]')  (Lxnen  program)  bx  the  .Alexander-xon- 
iiimiboldt  foundation  (to  BHS)  and  a  NSF  grant  EAR 
US().5ifi5  (to  Prof.  Karl  W.  Flessa.  Department  of  Ceo- 
seiences.  Unixt'isilx  o(  \ii/(iiia.  Tucson,  AZS5721).  We 
ari'  grateful  to  Kail  W.  i-1essa,  Daxid  L.  Dettman,  .An- 
drew S.  Cohen  and  i^axid  II.  (Joodxxiu  (Department  of 
CJeosciences,  Unixersitx  ol  Arizona.  Tucson.  ,\Z  S5721 ), 
and  jose  11.  Leal  for  helphil  discussions  and  to  txvo 
anonxinons  rcniexvers  who  improxed  the  manuscript. 
This  is  CT'IA.M  ((.'enlio  de  F.stndios  de  .Ahnejas  Muertast 
publication  no.  40.  The  authors  herexvilh  declare  that 
the  expel  iiiieiits  are  not  in  xiolaliiin  of  the  i-iirreiit  I  nit- 
ed  Stales  laws. 


LITKHA!  I  !;!•:  CflKD 

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THE  NAUTILUS  116(l):32-.35.  2002 


Paw  32 


Gone  with  the  wdnd:  a  pelagic  marine  species  described  as  an 
endemic  land  snail  from  the  Bahamas 


Philippe  Bouchet 

Museum  national  d'Histoiif  naturclle 

53  nil'  Buffon 

75()()5  Paris.  FKAXCF. 

pliouclictC'UiMilin.lr 


ABSTRACT 

AbaconUi  luiiijrui^d  (('Icntli,  19oSl.  hitliiTto  assuiiicd  tii  rcji- 
resent  a  monotvpical  genus  and  species  ot  Sul)nlinidae  (Gas- 
tropoda, Pulmonata)  endemic  to  the  Bahamas,  is  shown  to  l)e 
a  SMionvm  of  the  marine  pelagic  snail  Litiofxi  inehinostoma 
Rang,  1829  (Caenogastropoda,  Litiopidae).  Instances  of  genera 
of  land  and  freshwater  ga,stropods  originalK  mistakenly  de- 
scribed a,s  marine,  and  \ice  \ersa.  are  re\iewed.  Of  the  10 
names  invoK'ed,  four  are  \alid  (Daronia.  KiiroJns.  Tcivtropoiiui) 
or  potentialK  \alid  iDi'^onitixis).  one  (Bn^iulclhi)  has  been  sup- 
pressed In  the  K'ZN,  and  the  other  fi\e  iAiiiicoiiiil.  Afrocilli- 
klea.  Aiuiploidiniis.  Htilliiiujirrsiliii.  I'sciiihililnniiii)  are  s\no- 
nxnis. 


iXTHonrcTiox 

William  |.  CJench  (  19.>S)  ilcserihcd  Lcpllntinii  {Almciiii- 
ia  >  iHinfm'^u  as  a  new  .subgenus  and  spciics  in  the  piil- 
nionate  tauilK'  Siihiilinidae,  hasetl  on  s[)ec-inH'ii,s  eollei't- 
ecl  bvJ.C;.  and  V,.  C;reeii\va\- and  hinisclfin  the  Balianias 
in  April  1936.  The  clescription  was  aeeornpanicd  li\  hlaek 
and  white  photograph.s.  The  journal  wlieic  the  deseii[)- 
tion  \va,s  published  (Metnorias  de  la  Soeii'dad  ('ubaiia 
de  llistoria  Natural)  uia\'  ha\'e  contributed  to  the  lack 
ot  attention  that  these  new  taxa  ha\c  subsecjuentK  le- 
ceived  in  the  literature:  in  fact,  the  name  AlxicDiiia  is 
not  nientioned  at  all  bv  Zileli  (1959-fiO)  in  the  pulmo- 
nate  xohirne  of  the  standard  llaiidhucli  <lcr  I'dUiozoolo- 
ii,w.  Ahacoiiia  has  receiitK'  been  elevated  to  genus  lexcl 
by  .Schilevk.,  (1999:  513,'  fig.  666),  :uul  ehissified  :is  :i 
member  '<t  the  sublamiK  Subulininae. 

In  the  oii',:i!ial  publicalion,  C^^lench  noted  that  "this 
species  dil'i'i  r:.  :iuile  uol:ibK  from  all  others  known  to 
[hini|  in  this  genus  \1  rptinfiria].  The  remarkaliK  acute 
apex  and  the  :lKinge  in  the  sculpture  from  a.\ial  on  the 
first  four  \slioiis  ;.,  ,piral  on  the  remaining  whorls  is 
sharply  ihtfen-nt  IrMui  i,ther  species,  and  the  thiekeuiug 
of  the  shell  is  :ils.  "  'i-ual  hi  this  genus".  He  speculated 
that  Ij-plinrina  (A/.,.  <.■,■.  u(iufm<ia  anil  two  other  en- 
cleniic  pulmonates  mijjl  i.nstitute  a  reumaut  of  a 
much  older  fauna"  with  V\  .1  Induni  relationships.  How- 
ever, examination  of  the  type  materi;il  in  WV.'A  (Figiues 


1-3)  re\eals  tluit  Ahdiouid  lunifniiid  is  not  :i  snbidiuid. 
and  not  e\en  a  pulmonate,  but  :i  s\iion\  in  ol  IJIiopa 
iiirhiiuistoiiid.  d  circumtropic;il  pelagic  eerithioid  (Figs 
4-6).  (dencli  and  Schilexko  luiil  properK  described  tlie 
apical  sciilptiue  oi  Abdcoiiid.  but  had  tailed  to  recognize 
its  identibi.  The  protoconch  ol  Liliopd  incldii(>sl(iiiui  con- 
sists of  four  whorls  with  strong  a\ial  ribs,  aliiittiug  adax- 
ialK  on  the  preceding  whorl  anil  iiiti'rrupted  b\  a  smooth 
band  on  the  shoulder:  the  protoconcli/teleoconch  ilis- 
continuih'  is  nuirked  b\  a  sinusigera  notch  lF,iif|ue  et  ak. 
19SS:  l.SiZ.  figs  1,4). 

Liliopd  nu'ldnostddid  is  :iliiiiidant  in  (he  midst  ol  ll(i:it- 
ing  Sdr^d.ssiiDi.  and  it  is  eas\  to  imagine  how  tlie\  can 
be  waslieil  ashore  and  blown  inlanil  atti'r  the  se;iweed 
Iromls  base  been  sun-dried.  The  species  is  known  to 
occur  oft  the  15alKunas  (Liii|iie  et  ak,  19SS). 


.SYSTEMATIC  :.S 

Class  Ciastropoda  (  Jnier.  1797 
SiiperlamiK'  {Jerithioidea  l''leiniii'j:. 
FamiK  Litiopidae  Craw  1S47 
(Jeiiiis  Litiojid  Hang,  IS29 


1S22 


No\\  sxnoiiMii: 

Ahiicdiiiii  Clench.  19'1S.  T\pe  species:  I.cjUiiuirid  I Ah/ifimiri' 
iiidifniflii  (blench.  1938.  b\  original  desigiuition. 

Hoiiihijxiniis  lielanger  |iii  IjCssonj.  1835  is  another  ;ilreail\  rec- 
(ii.;iii/cd  s\ii()n\ni  o(  l.iliopii 

l.ilioj)!!  iiicldiii'sloNid  l^aiig.   1 S29 

\o"  sMionvni: 

I .(■jiliiiiiha  lAhiiioiiiai  luiufrii'^ii  Clench.  19.'i8:  321.  pi.  2-4. 

figs  1-2. 
Ahdconid  iiiinj)<iii<i — Scliile\ko.  1999:  514.  lig.  (i(i(i. 

Type  Ma(en:i1:      llolnlxpe  \IC'/  1  16705:  34  p:init\pes, 
\ic'/  116706:    I  p,n,il\pcs  NIC/  116707. 

Type  L<)c:ilil\:      S.uid  l^;iiik.  ( 'rossing  Baw  (ireat  .Abaco 

Isknid.  B:ili,iiii:is.  I'aralvpes  froni  the  l\pe  loc:ilit\  ( MC^Z 
Il6706):iiid  Mores  Island  (NIC/.  116707i. 

DISCUSSION 

Shells    of    kind    anil    Ireshwater    snails    :iie    occ:isioiKilK 
washed  to  llic  sea  :uid  ha\e  mil   ml  i  i-c  |iiiiill\    been  de- 


p.  Houclict.  2002 


Paffl"  33 


Figures  l-().  LiUdjiii  iiirhiiioslninii  Haiit;,  IS29,  1-3.  Il(il(it\pr  u{  Lipliiuiiiii  iAIkicoiiki  '  iKiuJni'^ii  Clciicli.  iy3S.  MCZ  116705, 
SiUKJ  Bank,  CJrossing  Bav,  Great  Abaco  Isiand.  Baliaiiias.  Heiijlit  4.2  nun.  4-(J.  One  (if  li\c  s\nt\pi's  fAlJtiopa  iiiclaiwsloma  Rang, 
1S29,  MNHN  [uncatalogued],  "Atlantic  Ocean".  Bang  coll.,  1S34.  Height  4.6  nnn. 


serihcd  as  marine  gastropiids.  Imt  the  rexcrse  lias  also 
occa.sidiialK  happened.  The  awareness  oi  tliese  sitnations 
aeross  tlie  tratlitional  boundaries  of  marine  and  non-ma- 
rine niollnscan  literature  is  problematic,  and  it  ma\  be 
uselnl  to  take  the  case  of  the  Abaconia  to  thaw  the  at- 
tention of  taxonomists  to  names  tliat  might  have  eseapeil 
their  attention.  These  are  presented  below  in  the  chro- 
nological order  tliex"  were  described,  ami  snmmari/ed  in 
Table  1.  Two  of  the  genera  invoKcd  e\en  became  the 
types  of  new  faniilv-group  names,  Tcri'troponiidae 
Hochebrnne,  1S81.  and  Anaplocamidae  I^all.  1921.  and 
it  is  probable  that,  if  tliex  had  not  been  mistaken  abont 
their  habitat,  Rochebnme  and  Dall  would  not  ha\e 
lailetl  to  recognize  them  as  representatixcs  ol  the  fami- 
lies Architectonicidae  and  Pleiu'oeeridae,  res]H'cti\cl\. 
Of  the  10  genera  inxoKed.  four  are  \alid  (l^arouui,  Kii- 
roliis.  Tcniwpoina)  or  potentiali\  \alid  (DiiSfutiuxis),  one 
(Bnmdchn)  has  been  suppressed  b\  the  K^Z.N,  and  the 


othei'  fi\e  iAlmci'iiid.  Alrocdiiiilcd.  Auiiplocdmii'i,  Bulli- 
uopiTsiliii.  I's<-ii(liilil)tniiti!  are  s\nou\ms. 

Daronia  A.  Adams,  1861.  Based  on  a  specimen  from 
the  Canning  ( .'olleetion.  CUfclostirmti  i  Daronia':  spinila 
A.  Aiiams,  ISttl,  was  described  as  a  new  subgenus  and 
species  of  niarine  gastropod  from  the  Philippines.  Waren 
and  l^oucliel  '  I9SS1  showed  that  it  is  a  South  .American 
species  of  the  land  snail  laniiK  l^oteriidae.  Daronia  \. 
.\danis,  IS()I.  being  a  senior  sxiioinni  lA'  Bncklcyia  Hig- 
'iius.  1872. 


Bromh'lia  iJoiirmiignat,  18(52.  l^escribed  as  a  new 
genus  of  \nc\lidae.  with  two  species.  B.  dronctiana 
Bourgnignat.  1S.54.  (originalK'  described  based  on  ma- 
terial from  the  ( fuming  collection  from  "North  Ameri- 
ca") and  B.  ^ihhosa  Bourgnignat.  1Sfi2.  said  to  be  li\ing 
on  damp  rocks  in  the  forest  of  Kdough.  near  Bone  in 
.Alsii'ria.  Brondclia  remained  in  the  freshwater  fainik' .An- 


l^uii'  34 


THE  NAUTILUS,  \'ol.  116,  No.  1 


Tabic  1.   XomiiKil  genera  of  marine  sjastropocls  iarranged  alphaliptically)  originally  described  a.s  h 


snail 


s  ana  Nice  \ersa. 


XoMiinal  "eniis 


I^escrilied  as 


Revised  position 


lieierenee 


Alxiroiiiii  C;lench,  193S 
Afrocanidca  Connolly,  1929 
Anuplocamus  Dall,  1S96 
Bwndchti  lionrgnignat.  1862 
Biillinopcrsilia  F.  N'ordsieck,  1972 
Daronia  .\.  Adams,  1861 
Di<loniaxis  Jousseaumc,  1889 
KiiroUis  de  Folin,  1870 
I'scudolilmnUi  Stcf'ani,  1870 
Teretropotna  Rochebrune,  1881 


Subnlinidae 

fresliwater  Bnccinidae 

Trichotropidae 

.\nc\lidae 

.\cteonidae 

Cyclostrematidae 

Fenissaciidae 

Acteonidae 

Daiidebaidiidae 

Cvclostomatidae 


Litiopidae 

Ranellidae 

Pleuroceridae 

Sipbonariidae 

FeiTJssaeiitlae 

Poteriidae 

P\Ta!nidellidae 

Fenissaciidae 

Pleunjbrancbidae 

.Arcbitectonicidae 


this  papt'i' 

Ben  (1998) 

Rehder (1942» 

Marsball  (1981) 

Smriglio  and  Mariottua  ,1996j 

Waren  and  Bonchet  (1988) 

Neubert  (1998) 

Zilch  (1959) 

Zilch  (1985) 

Crosse  ( 1882) 


cxlidae  until  Marsiiall  (1981)  recognizetl  it  as  identical 
with  tlic  marine  piilmonate  WdUamia  Monterosato. 
liS.S4  [famiK  Siphonariidae].  Incidentallv;  the  name 
Broiuh'lia  is  the  senior  s\nion\ni,  Init  it  \va,s  sub-seqiientlv 
suppressed  and  placed  on  the  Official  Index  ot  Rejected 
and  In\a!id  C.eneric  Names  in  Zoolog)'  h\  the  ICZN 
(1986.  Opinion  1410). 

Kciroliis  de  F"olin,  1<S7().  Kiinilii\  primus  de  FoHn, 
1870,  was  descrihetl  a.s  a  new  genus  and  species  of  ma- 
rine snail  trom  off  Vera  Caiiz  on  the  Pacific  coast  of 
C'entral  .Vnierica.  Kdrolw-:  was  not  originalK  placed  in  a 
hiTuiK;  and  was  omitted  Irom  standard  textbooks  and 
treatises  lor  neark  a  centun.  It  was  treated  bv  Zilch 
(1959)  as  a  \alid  subgenus  ol  Ccrilioidcs  Fenissac,  1S14 
[laTuily  FerussaciidaeJ,  and  h\  Schileyko  (1999)  as  a  lull 
genus. 

Psc'tidolihduia  Slefani,  1S79.  Daiidchardia  larcidina 
•Stelam  and  I'antanelli.  1879,  was  described  as  a  land 
snail  Irom  (he  hills  near  Tarantn  in  southern  Italy.  It  was 
made  tiu'  t\]ie  species  of  a  section  Psriidoldiania  Stefani, 
1879,  ol  Daiidchardia  Ilai-tmann.  1821  |lamiK  Dande- 
bardiidae],  later  elevated  to  full  genus  b\  Forcart  (1950). 
Zilcii  (1985)  has  shown  that  D.  larcittiiia  is  a  svnonvm 
ol  Siisania  tesliidiuaria  Cantraine,  1835,  and  I'srudoli- 
hania  a  .synonym  n\'  Sii.saiiia  Cray.  1857  |laniil\  Fleuio- 
brancliidae]. 

Jeretropnma  Rochebrune,  I8S1.  Inunediatc  l\  :ift(  r 
die  description,  based  on  live-taken  specimens  fiuui  the 
Dakar  peninsula  (Senegal),  of  Tcirlropoiiw  pcrricri 
Kocheb-rmii'.  1881,  as  a  new  genus  and  species  of 'Cv- 
clostomace.i'.  and  type  of  a  new  subfamiK- Terelropom- 
idae  (sic  Cross.'  1SS2)  recognized  it  as  an  architecton- 
icid  relalet!  to  lonuia  Cray.  1842.  Tercimpoma  is  cur- 
rently treated  as  a  \<i!id  subgenus  of  Heliactis  d'Orbignv, 
1842,  and  pi-rricn  is  rcg-arded  as  an  .Mlantic  subspecies 
of //.  infiiiidiliidif.)ri!ii\  fCmclin,  1791)  (Bieler,  1993). 

Dificminxin  Jousseamnr.  1889.  Di<i(>iiiaxis  houroui<i- 
nati  Jousseaumc,  is^',),  ,\as  described  as  a  new  genus 
and  species  from  vi.  n  .uid  trt\ited  a.s  a  land  snail  of  the 
family  FenissacliiL;-  b\  ifs  author  and  bv  Zilch  (1959: 
342).  BiLsed  on  a  re-exaniinalion  of  the  t\pe  material. 
Neubert  (1998)  realloialed  it  to  the  marine  famiK  l'\r- 


amidellidae.  Although  Schile\ko  (1999),  relering  to  Neu- 
bert, suggested  that  onl\  an  anatomical  stuck  would  es- 
tablish firmk-  the  systematic  jxxsition  of  D/gofuV/.v/s,  the 
heterostrophic  protoconch  lea\es  no  doubt  tliat  it  is  in- 
deed a  p\  ramidelloid. 

Anaplocamiis  Dall,  1896.  Anaplocamtis  horealis 
Dall,  1896,  was  described  as  a  new  genus  and  species  ot 
Trichotropidae  from  .Alaska,  and  later  e\en  became  the 
t\pe  of  the  monotvpic  famik  -\naplocamiclae  D;ill,  1921. 
Rehder  ( 1942)  discoxered  that  this  was  in  fact  a  SMionvm 
of  the  freshwater  snail  Aiiciilosa  ddatata  Conrad,  1835 
[now  Lcptoxis  dilatata:  famiK'  Pleuroceridae],  from  east- 
ern United  States,  which  had  been  erroneously  labelled 
as  from  a  marine  .Alaska  localitw 

Afrociniidea  Connolly,  1929.  Afrocanidca  ^^cmma 
C'onnolk.  1929.  was  desciibed  as  a  new  genus  and  spe- 
cies of  freshwater  Bnccinidae,  reputedk'  Irom  Shimbi 
Hills,  Kenya,  but  the  holot\pe  was  shown  to  be  a  lanal 
shell  of  Cipuatiiiin  imtriciniuii  (Roding.  1798)  [family 
Ranellidae],  with  Afrocanidca  Connolk,  1929,  ending  up 
in  s\"n()u\nn- of  Cw(^//mi(»//(  Mi'irch,  1852  (Beu,  1998  i. 

BullinopersiUa  F.  Nordsieck,  1972.  Based  on  a  sin- 
gle empt\    shell,   BullinopersiUa  sjihacroidcs   F.   Nord- 


.■k,  197: 


was  itescritied  as  a  new  genus  and  species  ot 


.\cteonidae  from  shallow  water  off  the  Baleares,  in  the 
westeiii  Mediterranean.  Smriglio  and  Mariottini  (1996) 
recogni/ed  it  as  a  jii\iaiik'  of  the  land  siuiil  Fentssacia 
folliculus  (Clmelin,  1791'  [lamily  FeiTissaciidae]. 

IdTER.ATURK  CITFD 

Ren,  A.  1998.  Indo-West  Pacific  Ranellidae.  Bnrsidae  and  Per- 
sonidae  (Mollnsca:  (Jastropoda).  A  monograph  of  the  New 
(Caledonian  l.inn.i  and  revisions  of  relatetl  t;L\a.  Memoires 
dn  \hisenni  national  d'llistoire  naturelle  178:  1-255. 

Bieler.  H.  199.'V  Aidiiteetonicidae  of  the  Indo-Pacific  (Mollns- 
ca. (iastropod.i).  .Vbbandlungen  des  Natnnvissenseliaf- 
tlielien  N'ereins  in  Ilanibnrg.  new  series,  30:  1-.377. 

C'lencli.  W.  |.  I93S.  Land  and  Iresliwater  mollusks  ol  Ciand 
Bahama  and  the  .\baco  Islands.  Bahama  Islands,  Memo- 
rias  de  la  .Sociedad  (.iili.ui.i  de  llistiiri:i  Nalnral  12(4): 
.3()3-,3.33.  pi.  24-25. 

("rosse.   11.    ISS2.  [Book  review   ol|  Sin   mi  Ivpe  nouvian  de  l;i 


p.  Boucliet.  2()(»2 


Wm-  35 


faniille  des  C'vclostoniaoea,  par  le  Dr.  A.T,  dc  Koclicliniiir 
|ournal  de  tA)iich\liolosjie  30:  249-250, 

F'liriart.  1.  1950.  S\steniatique  dcs  MolliiscjUfs  en  Idiiiic  Ai' 
Duudcbardia  et  re\ision  des  cspetx-s  d'.'Viuitolic  ct  dc  I'ilc 
de  Crete.  Journal  de  Conchvliologie  90:  1 07-1 17. 

Luque,  A.  A.,  Templado,  ].  and  L.  P.  BiirnaN'.  HISS.  On  tlic 
systematic  position  of  the  p;enera  Litiopii  Rans;.  1829  and 
.\lilhii  II.  and  .\.  ,\danis.  1S.5.).  Malarol(ii;i(;il  Ri'\ii'\\, 
suppl,  4:   I  SO- 19.3. 

Nhnsliali,  B.  A.  1981.  The  ij;enns  Williaiiiia  in  llir  ucstrni  Pa- 
rific  (.Mollusca;  Siphonariidae).  New  ZciI.likI  |()iinial  ol 
Zoologv-  8:  487^92. 

Nenbert,  E.  1998.  Digoniaxi.s  h(>nri;,iiii:,iiiiti  |ou.ssfaunie  1S89, 
a  genus  and  species  of  the  Pvraiiiidellidac  ( tlastropoda; 
Heterostropha).  Archiv  Fiir  Moliuskcnknnde  127(1-2): 
10.3-106. 

KIZX  [International  Commission  on  Zoolo'^ical  Nomencla- 
ture]. 1986.  Opinion  1410.  W'iUUimUi  .Monterosato,  1884 
(Mollusca,  Gastropoda':  consencd  Biillrtiii  ol  Zoolo<4ical 
Nomenclature  43:  249. 

Rauij,  S.   1829.  Notice  sur  le  Litiopc,  Litiopa.  iji'un-  ii(iu\cau 


(Ic  Molhisque  gasteropode.  .\nnales  des  .Sciences  Natu- 

rcllcs  Hi:  .30.3-.307. 
liilidii    II     \    1942.  .\  note  on  tin-  'icmis  AiKiplocniiiiis  l.)all. 

I'll.'  Naiililus  .56:  49-.5(). 
Schilc\ko.  A.  .\.  1999.  'IVcalise  on  Hcccul  terrestrial  pulinonate 

molluscs.  Part  4.  Kutlicnica  suppl.  2:  4.35-564. 
Snuiglio.  C.  and  I',  .Mariottiui.   1996.  Central  'IXTrlieuian  Sea 

.Mollusca:  Xi.  Description  oiCiillosinicon  ti/rrlicnicnin  sp. 

uo\.  (Castropoda,  Acteonidae)  and  remarks  on  the  other 

.Mediterranean     species     ol     the     lamilv     .Vileonldae 

d'Orhi^nv.  18.35.  Basteria  60:  lS.3-193. 
W'an'n.   \   and  1'.  Bouchet.  1988,  .\  new  species  of  X'anikoridae 

Ircini   the  western   Meiliteiiauean.  with   remarks  on  the 

northeast  .Atlantic  species  ol  the  taniik.  Bollettino  Mahi- 

ci)lo<iico  24(.5-8):  7.3-100. 
Zilch.  A.  19.59-1960.  Teil  2:  F.uthvncina   In:  Oil.  Schiutlcwoir 

led.  I.  Hiiiullmch  tier  PiiliiozooUi^c.  Band  6,  Caslnipudd. 

Bomtraeger.  Berlin,  \ii  +  8.35  pp. 
Zilch,  .\.  19S5.  Ddudflxirditi  ttiri'iitiua  und  I'sciuhilibcniid  i.Nu- 

dihranchi:i:    N'otaspidea:    Pknirohranchidae.    .Archix    liir 

Mcilluskvnknndc  115i4-6.:  291-.30I). 


In  Mciiioriaiit 

Tiulaslii<2;t'  Hahe  Rus,sel  H.  jciiscn 

1916-2001  191S-2()01 


TIIK  NATTILUS  116(l):36-38.  2002 


Paw-  36 


Note 


Biogeogniplu"  ol  the  deep-sea  gastropod  Palazzia  plaiiorbis 
(Dall,  1927):  an  uncommon  form  of  rarity 

Michael  A.  Hex 

Departiiient  ol  Biolog\' 
Unix'cisih'  of  Miissacliiisptts 
100  Mon-isse\  Blvd. 
Boston,  MA  ()21iT  USA 
iriiclia<'l.ro\(a'\iiiil).e(lii 


Palazziii  plaunrhis  iDall,  1927)  is  a  iiiiniitf  dri'p-sea  gas- 
tropod ol  uncertain  plniogenetic  atfinitv,  possibly  resid- 
ing in  one  ot  die  groups  that  were  once  included  in  the 
Archaeogastropoda  (W'aren  1991,  1993),  It  has  a  plani- 
spiral  shell  hearing  distinctive  iixial  ribs  that  branch  mid- 
\va\^  towai'd  the  outer  whorl  margins  (Figure  1),  Like 
many  deep-sea  species,  its  natural  liiston  is  unknown, 
.Vloore  (1971)  first  noted  that  P.  plaiiorliis  occupies  an 
innisnalK  broad  geographic  distribution  in  the  western 
North  .Vtlantic,  lia\ing  been  dredged  at  2967  ni  oft  Lali- 
rador  and  at  53S  ni  oil'  Florida,  W'aren  ( 1991,  1993)  sub- 
sequently reported  the  species  \mu\  west  of  Iceland  at 
241  111  and  in  the  eastern  North  .Atlantic  o IT  the  British 
Isles  Iroin  2081-2898  in.  Here.  1  proxide  additional  re- 
cords lor  tlie  western  and  eastern  North  Atlantic,  tidp- 
ical  Atlantic,  and  the  western  South  .Atlantic  (Tabli'  1). 
These  new  localitv  data  come  Irom  m\'  examination  of 
material  pro\id(>d  In  the  Woods  Hole  Benthic  .Sampling 
Program  (Sanders.  1977)  and  tlie  .\tlantic  Continental 
Slope  and  Rise  Stuck-  (Cirassle  and  Maciolek,  1992).  Pal- 
azzia planorbis  is  revcaleil  to  have  an  e.xtraordinariK 
broad  geographic  distribution  that  spans  Southern  and 
Northern  Hemispheres  IVoin  43°33'S  to  (i5"ori'N,  and 
includes  both  the  eastern  and  western  corridois  ol  the 
deep  .Atlantic  (Table  I,  Figure  2),  Its  bath\  nietrit-  range 
is  similarl)  impressive,  extending  IVoni  the  uppermost 
reaches  of  the  continental  slope  (241  m)  to  the  deep 
alnssal  plain    521fi  m). 

While  the  shell  sculpture  of  Palazzia  planorbis  ap- 
pears to  be  niiiijue  among  [)lauispiral  deep-sea  micro- 
gastropods,  tliere  is  some  individual  variation  in  the  ex- 
pression and  nnmbcr  of  axial  ribs  and  the  extent  of 
branching.  Ihr  p.iltern  of  geographic  variation  throngh- 
oiit  its  ranwe  is  ditlien'L  to  quantifV  becaii.se  of  the  scar- 
city ol  material  inu  wid  ■  separation  of  sampling  locali- 
ties (Table  1.  i'lenn  2  However,  the  specimen  from 
the  eastern  tropica!  At!  nitu  shown  in  Figure  1  is  re- 
markably similar  in  uiik  at.i.Kc  to  those  fimired  bv  War- 
en  (1991:  77,  fig,  ih  .\-lji  iliat  were  collected  in  the 
we.stern  and  eastern  Nurih  Atlanli  913  to  3259  m  ,slial- 
lower  and  45°  to  70°  t(    ihe  n.nlh. 

Its  enormous  geograpliic  and  ballivnietric  ranges  are 


espeeiallv  notable  because  Palazzia  jAanorhis  is  vei"v 
rare,  and  is  included  among  the  smallest  gastropods. 
Specimens  from  the  Woods  Hole  Benthic  Sampling  Pro- 
gram (Table  1)  were  all  collected  bv  using  epibenthic 
sleds  (Hessler  and  Sanders,  1967),  which  are  txpicallv 
tovvetl  for  (jue  kilometer  on  the  bottom  and  vield  sam- 
ples that  represent  about  HHIO  linear  nr  of  seafloor.  As 
a  rough  approximation,  the  data  in  Table  1  suggest  a 
median  densitv  of  one  individual  per  1000  nr,  with  a 
maximum  of  17  individuals  per  1000  m-  in  the  western 
South  Atlantic.  This  is  rare,  even  for  deep-sea  gastropods 
which  usuallv  exist  at  low  abundance  (Rex  et  al.,  1990; 
Rex  and  Etter.  1998).  With  a  shell  diameter  reaching 
onlv  between  one  and  txvo  inillimcti'rs,  it  is  also  an  ex- 
tremely small  gastropod — bolli  lor  the  (Gastropoda  as  a 
vv'hole  (Bieler  and  Mikkelseii,  1998)  and  tor  the  dee^p- 
sea  fauna  (Rex  and  Etter,  1998), 

Ri'latioiishijis  among  geographic  range  size,  abun- 
dance and  both  size  are  central  to  macroecologV'.  Al- 
though a  grt'at  ileal  ol  variation  has  been  obseived, 
abundance  and  bodv  size  are  generallv  related  to  one 
another  negativclv,  and  related  to  range  size  positivelv 
(Caston,  J994;  Brown,  1995;  Gaston  and  Bhickbimi, 
2000),  Thus  the  coinbiiKition  ol  low  densitv,  large  geo- 
graphic nmge  and  small  bodv  si/e  obsciAed  in  Palazzia 
])l(iiu)rl)is  is  exceptional.  Biibinowitz  i,  I98|l  proposed  ;i 
threc-w;iv  (.-kissifieiition  ol  raritv  based  on  geographic  dis- 
tiibntiiin,  habit;it  speeificitx  and  local  population  size. 
Palazzia  planorbis.  with  its  extensive  geographic  range 
in  the  Atlantic,  oceup;ition  ol  dillerent  li;ibitats  ideptli 
zones)  and  small  inral  populations,  represents  one  ol  till' 
most  muonmioii  lorms  ol  raiitv  in  the  Ivabiiiowitz 
scheme.  Schoenei'  (1987'  releried  to  low  ii'hilive  ;ibnn- 
(kmce  throughout  a  s|H'cii"s  rangi'  as  siillusive  raritv.  and 
showed  (hat  this  p;illeni  is  an  extremi'  euil-membi'r  ol 
the  r;iritv  coiilinimiii. 

\\  idelv  distributed  rare  spi'cies  are  also  known  among 
dei'p-se;i  protobranch  bivalves.  Examples  include  Pris- 
liillouw  iiitois  and  /'  alba  (Sanders  and  .Alien,  1973),  :ind 
Yiildirlla  fabula  (Allen  et  al.,  1995).  Other  bivalves  are 
connnon  in  one  or  moi'e  deep-sea  b:isins  and  rare  else- 
where (.Allen  .ind  Sanders.    U)96).  a  pallcin  that  Schoe- 


M.  A.  Hc\.  2002 


Paec 


Fiijiiro  I.  I'lilm-iii  plunoiiiis  collected  Iidiii  station  U)5  m 
the  casttTii  tropieal  Atlantic  (si'l-  Talilf  1  Im  l(i(alit\  data'  Tin- 
specimen  measures  1.24  mm  in  ina\niiuni  (lianielei. 


nei'  llVJ.ST)  teniR'tl  tlittusi^e  rafitx.  'Hie  iiiciili'iiee  ol  lare 
deep-sea  gastropods  oceHp\iii'4  ludad  geographic  ranges 
is  still  unclear  since  most  taxononiic  s\ntlieses,  siicli  as 
Boncliet  and  W'arens  (19S0)  excellent  monograph,  haxc 
been  regional  in  scope. 

Studies  of  raiitx'  and  I'ange  delineation  are   liaiiglit 
with  imt'ertaiiit\  and  methodolosrical  diificnlties  (Ciaston 


U)U  I  It  should  he  eaiitioned  that  the  apparent  aliun- 
danee  ol  I'dldz-Jd  j}ltiiiorl}i\  iTahle  1)  nia\"  be  paitK"  due 
to  sampling  error.  The  mesh  size  of  the  collecting  net 
used  on  epibenthic  sleds  deploxed  in  the  Woods  Mole 
Beiithie  ,Sam|)ling  Program  is  1,(1  mm.  Hiis  is  fine 
enough  to  e.iplure  adults.  Smaller  jn\enil<'s  could  pass 
through  the  mesh,  but  some  do  in  lact  appear  in  sam- 
pk's.  The  siexc  si/e  used  to  sort  the  niacroiauna  is  420 
fjLm.  whii'h  should  retain  .ill  but  the  smallest  post-lanal 
iudi\idiials  ol  this  spi'cies.  Karlier  sampling.  lio\ve\'er, 
used  larger  mesh  and  sii'Xi'  sizes,  suggesting  that  rec'0\'- 
er\  of  /'  pliiiiorhis  was  happenstance.  .Since  the  samples 
aie  widi'K  separated  and  lepiesent  onl\  fi\e  regions  of 
the  Atlantic  I'lable  1,  Figur(>  2),  we  cannot  doeiiinent 
the  full  geographic  range  or  know  relatisc  abundance 
th.roughout  the  range.  It  is  possible  that  R  pUiiiorhis  has 
an  unrecognized  ceiilei'  of  high  ileiisitx'  and  shows  dif- 
fusi\c  lather  than  suffusixc  raiitx"  (sensw  Schoener. 
19S7).  l'"iuall\.  it  should  be  pointed  out  that,  while  V. 
phiiioiiiis  appears  tii  be  a  coherent  morjiliospecies.  some 
pheuot\]iicalI\  unifoim  deep-sea  mollusks  show  strong 
population  structure  at  the  genetic  level  (Etter  et  ah, 
19991,  /'  jiliiiinrhis  mas  be  a  coiuplex  of  geneticallv  di- 
\ergent  populations  or  species.  E\cn  considering  these 
potential  probleiiis,  the  combination  of  a  Pan-.Atlantic 
distribution,  consencd  moiplioiogw  low  abundance  and 
siriall  size  would  seeiu  to  represent  an  extremeK'  unusual 
form  of  raritx  lor  lu.niue  mollusks. 


Table  1.  Localitv'  data  for  Palazzia  plaii(>ii)i\  in  the  ,\tlantic.  Where  the  source  of  data  is  not  indicated,  it  refers  to  this  study. 
.XCSAK  refers  to  the  Atlantic  Continental  Slope  and  Hise  StuiK  (Grassle  and  .Maeiolek.  f9!)2i.  WliOl  refers  to  the  Woods  Hole 
Benthie  Sampling  Program  (Sanders.  19771.  and  is  followed  In  the  name  of  the  \essel  and  iruisi-  immher.  Please  see  references 
listed  under  source  for  information  on  other  ((illcelm^  exprditions 


Kegion 

Source 

lv\pe(fitiou 

Station 

l)e[)thilll 

F.lUdule 

Fuiigitude 

Specimens 

West, 

111  North  .\tlaiitie 

Wareii. 

]m.\ 

|(ni  lidgason.  ( .'oil. 

— 

241 

fi.5°0fi'  N 

26°42'  W 

Moore. 

1971 

\U  1  1,1)1  ic 

— 

20(i7 

55°3fi'  N 

54°33'  W 

WllOI/CllMN    fOf-i 

334 

4400 

4()°43.3'  \ 

46°14.2'  W 

6 

ACSAK 

13 

12.50 

39°48.35'  N 

70°  .54.94'  W 

WIIOl/Aii.WTis  II 

24 

12(S 

:5S0(i 

39°37.25'  N 

66°45.55'  W 

\\ll()l,/K\oHH  .3.5 

.34(1 

.3310 

.3S°l(i.O'  N 

70°21.55  W 

WIlOf/AirwTis  11 

to 

17.5 

KiSO 

.36°.36.()'  N 

fiS°3()'  W 

Moore. 

197f 

.\l,H\rHciss 

26riS 

.5:5S 

.3(f5S'  N 

79°3S'  W 

Faste 

rn  North  ,\tlaiitie 

Waren. 

f99.3 

INCAI, 

DSOl 

2001 

.57°59.5'  N 

1()°40.5'  W 

Waren. 

199.3 

INCAF 

I)S()2 

20S1 

.57°58.7'  N 

10°48.9'  W 

\\"aren. 

1993 

INCAF 

nS(l5 

2.503 

.5(i°27.9'  N 

ini.s' W 

Waren, 

1993 

INCAI. 

DSOti 

2494 

.56°26.3'  N 

irio.fi'  w 

Waren. 

1991 

INCAF 

CPOo 

2SS4 

.5.5°0().4'  N 

12°30.2'  W 

Waren. 

1993 

Cn,\i.i,i:N(a  n  II 
Wll01/f:ii\i\  10(1 

190 
:52,'5 

2  SOS 
.3:547 

.54°4F  N 
.5()°()S.3'  N 

12°1S'  W 
|.3°.52.3'  W 

W.'sti 

irii   lYopieal  ,\tlaiitie 

W  ll()i,K\oKH  15 

295 

1011 

S°()4.2'  N 

.54=2 F3'  W 

W  IIOl/.Vri.WTis  II 

31 

1.50 

SS7 

7"5S.()'  S 

.34°22.0'  W 

Fast( 

•rii  Tropital  .\tfiiitic 

WIIOI/.Vri.WTis  II 

42 

lOS 

45(s:5 

I(r26.5'  S 

9°()6.5'  F 

\\II(M/.\tl\\tis  II 

42 

197 

459tt 

10"29'  S 

9°04'  E 

WllOI/.Vri WHS  II 

42 

195 

3797 

14^45'  S 

9°55'  E 

West. 

erii  South  .\tlaiitic 

WIlOI/Ari.WTis  II  fit) 

239 

l(i70 

.36°49.()'  S 

.5.3°  15.4'  W 

WII()l/.\Ti WHS  II  (ill 

247 

.52  Ki 

4.3°33.()'  S 

4S°5S.l'  W 

17 

Pa<ie  3S 


THE  NAUTILUS,  Nol.  116,  \, 


Figure  2.  I'lic  (listrilnitiiin  ol  P(il<r.-Jii  jihnutrhis  in  llic  .'\t- 
liUitif  (.sec  'l"al)le  1  lor  stiitioii  data).  Oirclfd  nuiiibeis  represe-nt 
the  niinihcr  ol  samples  xieldinij  P  pldiiorhis  in  that  region. 
( .'ircles  wiliioul  nuiiiliers  represent  iiiilividual  samples. 


.'\fK\0\\I,Kn(:\IF,\TS 

I  thank  .\iulrea  iiex,  Hon  Kttcr,  .\nders  W'aivn.  Pliiiippe 
iioiichi't,  and  ('arol  Stuart  lor  llieir  eoniiiicnts  on  tiiis 
paper,  jack  ('ook  lor  the  map  of  the  Xtlantie  and  Maria 
r'apiisja  lor  iielpintj  to  prepare  the  manuscript.  The  ma- 
terial from  the  Woods  Hole  Bcnthic  Sampliuii  Pronrani 
wa.s  provided  by  Howard  Sanders,  and  From  the  .-\tlantii- 
Continental  Slope  and  Hise  Stud\  In  James  Blake.  This 
research  \va.s  .supported  1)\  the  National  Seienee  l-'oun- 
dalion  CIrant  C)(  :i':-9:3()l(iS7  and  In  llie  Uni\eisit\  ol' 
.Vlas.saclinsetts. 

iJTEIl.VrUHE  CITED 

•\1!(  n.  J.  ,\    and  II.   L.  Sanders.  1996.  The  zootjeoijraphv.  di- 
M'liii'.    liici  orinin  ol'  the  deep-sea  protobraneh  hixakes  ol 


the  Atlantic:  The  epilo^ne.  Progress  in  Oceanography  .38: 
95-15.3. 

Allen.  J,  A.,  H.  L.  Sanders  and  F.  J.  Hannah.  1995.  Studies  on 
deep-sea  Protohranchia  (BixaKiaJ:  the  subfamily  Yoldiel- 
linae.  Bulletin  of  the  British  .Museum  (Natural  Histor\), 
Zoolog;)  61:  11-90. 

Bieler,  R.  and  P.  M.  Mikkelsen.  199S.  Aiiiinoniccni  in  Florida: 
notes  on  the  smallest  li\ing  gastropotl  in  the  United  States 
and  comments  on  other  species  of  Omalogxridae  i  Het- 
erobranchia).  The  Nautilus  111:  1-12. 

Bouchet,  P.  and  A.  VV'aren.  19S().  Rexision  of  the  north-east 
,'^tlantic  bathyal  and  abxssal  Turridae  (Mollusca,  Gastro- 
poda). The  |()\n"nal  ol  Mollusean  Studies.  Supplement  S: 
1-119. 

Brown,  |.  H.  1995.  Macroi'cologN.  Uni\(Msit\()f  Chicago  Press, 
( 'hicago,  269pp. 

Ftter  R.  ]!,  M.  A.  Rex,  M.  C.  Chase  and  J.  W.  Quattro.  1999. 
A  genetic  dimension  to  deep-sea  biodi\ersitx.  Deep-Sea 
Research  I  46:  1095-1099. 

{".uston.  K.   |.   1994.  Raritv.  C:liapman  and  Hall,  Foiulon.  205 

PP 
Caston.  K.  J.  and  T.  .\1.  Blackburn.  2000.  Pattern  and  process 

in  macroecolog\.  Blaek^well  Science.  Oxiord.  377  pp. 

(irassle,  J.  F.  and  N.  J.  Maciolek.  1992.  Deep-sea  species  rich- 
ness: regional  and  local  di\ersitx'  estimates  Irom  quanti- 
tative bottom  samples.  American  Natundist  139:  313-341. 

Hessler,  R.  R.  and  H.  L.  Sanders.  1967.  Faunal  di\ersit\'  in  the 
deep-sea.  Deep-Sea  Research  14:  65-7S. 

Moore,  D.  R.  1971.  .\  deep  water  Omtiltnj^ijid  in  the  wt'stern 
.'Vtlantic.  The  Nautilus  84:  113-117. 

Rabinowitz,  D.  1981.  Se\eu  forms  of  raritv.  hi:  Svnge,  H.  (ed.) 
The  biological  aspects  ol  rare  plant  consenation.  pp.  205- 
217.  Wiley  New  Y<n-k. 

Rex,  M.  A.  and  R.  J.  Etter  1998.  Bathsnietric  patterns  ol  bod\ 
.size:  implications  for  deep-sea  biodi\ersit\'.  Deep-Sea  Re- 
search II  45:  10.3-127. 

Rex,  M.  A.,  R.  J.  Etter  and  P  W,  \nneskern.  jr  1990.  Densitx 
estimates  lor  deep-sea  giistnipotl  ,issenibl,i<4es.  Deep-Sea 
Research  37:  .5.55-569, 

Sanders,  II.  L.  1977.  K\c)lntion:in  eeolog\  :md  the  deep-sea 
benthos.  l\i  (ionlden,  (.'.E.  led.  i  The  changing  scenes  in 
natural  scienics  1776-1976.  pp.  22.3-243.  .-Kcadenn  ol 
N:itnral  Sciences  Special  I'nblication,  Philadelphia. 

Sautlers.  II.  L.  and  |.  .A.  .Vllen.  1973.  Studies  on  the  deep-sea 
Protobrancliia  (Bixakia);  Prologue  and  the  Pristiglomitlac. 
Bulletin  ol  the  Museum  ol  (^omparatixc  /oologs  145: 
237-262. 

Sthoenei'.  T.  W.  1987.  The  geogniphical  distribution  ol  rarit\. 
Oecologia  74:    161-173. 

Wan'ii,  A.  1991.  New  and  little  known  Mollusca  Ironi  Irel.md 
.md  Scandinavia.  Sarsia  76:  5.3-124. 

\\:in  II  \.  1993.  New  and  little  known  Mollusca  liciiii  Icej.md 
.md  Scandina\i;i.  P:irl  2.  Sansui  78:  1.59-201. 


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THE  NAUTILUS 


Volume  lib.  Number 2 
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ISSN  0028-1344 

A  quarterly  devoted 
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vv  joa;  rfi'.-i  Ocfe--r.og(spl;ic  inshiution 


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CONTENTS 


\'(>hiiiif  IK).  Xmnhcr  2 

June  IH.  2002 

IS'SX  002H-1344 


Russell  L.  Million  A  cladistic  anaKsis  ol  LUha\'ui  i  (lastnipoila:  I'li-uiDtrridae) 

iisiiiy;  inoipliold^ical  c'liaraftci's  39 

Kic-ardo  Ne«a  i'itrtilliojilida  krinfilonniL  a  new  species  l(  iasti'opoda: 

Hodollo  ^<-'l^a  ( 'oialliopliilitlae)  Iniin  tlie  (Jaiian'  Islamls  li\iii'4  (in 

Ajigel  A.  Luque  Aulipallics  in>ll(i\li>iii  (( 'nidaria:  Antlio/na:    \iili]iatliaiia! 50 

Martin  A\er\  Sn%der  I'ushiiis  iloi  jxh-di.  a  new  spi'cies  ((iastrnpoda: 

I'aseidlaiiitl.ie)  liiiin  tin-  Ri'd  Sea.  and  lan^e  e\lensi(in  lor 

two  odier  species 56 

Helena  Fortunato  Tlie  systematic  position  ol  Sln>iiiliiiiu  \('(il<>ii(ij)sisi  liiidiic 

Petncli.   I9SS  (Castropotla:  f.'olniiiliellidae) 59 

H.  Lee  Fairbanks  The  repiddncli\e  aiiatonix,  t;L\ononiic  slatns,  and  raii^e  ol 

Oivolu'lix  (iljiiiKi  (I'drotl.  19011  iCiastropoda:  I'ninionala: 
Oreolielicidae! 62 

Notes 

Bruce  A.  Marshall  Antliorsliip  and  ilate  of  |inlilication  nf  (hirca  r7///rii\/,s 

I'liilippi  /■)(  Kiister.  1S44  (Bi\aKia:  Ostreidae) 66 

Paul  Callomon  Tadaslii'^e  ilal.e  1 1916-2001 ) 67 

Book  re\  iew 6J 


THE  NAUTILUS  116(2):39-49.  2002 


Paee  39 


A  cladistic  anal\  sis  of  Lithasiii  (Gastropoda:  Pleuroceridae)  using 
moiphological  characters 


Russell  L.  Minion' 

IJcparlMiciit  ol  Bu)l()j;ii'al  Siiciict's 
Uiii\ersih-  ol'  Alahanui 
Tuscaloosa.  AI.354S7  USA 


ABSTRACT 

Till'  fla.ssificatinii  ol  plciiroiinil  snails  ami  otlirr  Ircsliwatcr 
iiiolhisks  has  liistoricalK  liccii  hascd  on  inoipholo<;ic'al  char- 
acters. Dcspiti-  \cars  ol  ta\oiioiiiic  work  on  plenroccrids.  no 
sinijlc  work  inchides  all  rccm^ni/ed  ta\a  Ironi  a  u;i\cn  tjronp 
anil  onl\  a  tew  systematic  treatments  ol  the  lamiK'  or  indixidnal 
genera  exist.  Modi-rn  methods  ol  ph\  loijenetic  s\stematics  ha\c 
shown  that  some  moipholoi;ical  traits  do  not  support  liistori- 
calK accepted  mollusk  classihcations.  II  anaKses  ol  nioqiho- 
logical  characters  do  snppoil  ciuix-nt  taxononiic  li\potheses, 
then  the  classification  ol  tliesi-  e;roups  can  lie  tonsidered  as 
stable.  It  not.  our  approach  retjarding  diagnostic  cli.u'acters  tor 
these  groups  must  change.  This  paper  uses  25  shell  and  radular 
characters  in  a  cladistic  anaKsis  ol  the  pleurocerid  genus  Ijth- 
asiu,  and  compares  flie  findings  to  prexionsK  snggestt'd  clas- 
sifications ol  the  group.  Cladistic  anai\ses  do  not  support  an\ 
current  or  historical  classification  ol  Litliiisiii.  However,  these 
moiphological  characters  are  found  to  rongliK  delineate  IJth- 
iisid  and  other  extant  pleurocerid  genera,  challenging  pre\ious 
works  that  suggest  such  characters  ha\e  limited  ntilitx. 


IXTKODUCTIOX 

HistoricalK.  the  i-lassific-atinii  ot  tresliwater  iiKilhisks  has 
relied  lieaxiK  on  iiidiphdldgical  features  such  as  shell, 
soft  anatoiiu',  and  reproiluetive  .structures  (e.g..  Tnou. 
LS73:  Heard  and  Cuekert.  1970:  Davis  and  Fuiler.  19S1; 
Burcli  and  Tottenham.  19S()).  Thks  is  partieiilarK  cxident 
for  pleurocerid  snaiks,  win-re  sliell  features  aec-ount  tor 
the  inajorit\'  of  diagnostii-  characters  used  in  the  taxon- 
omy and  clas.sifieation  ot  the  group.  One  grouji  ol  ph-ii- 
roeerids  that  lias  !)eeii  clas.sified  on  the  liasis  of  shell 
eharacti-rs  is  Lilliusid  llaldeiiian.  IS40.  a  genus  ol  large 
ri\'er  snails  loiind  tlirongliout  tlic  ( 'nnilierland,  Ohio. 
Mi.ssissippi,  and  Tennessee  River  drainages. 

Species  of  Lilliasia  pos.sess  eonie  to  o\ate-conic  shells 
with  tusiforin  apertures,  a  posterior  eallns  on  the  parietal 
wall,  and  fre(|neiitl\  some  degree  ot  sculpture  on  the 
hoiK  whorl  (linrcli.  19'S2).  .Most  species  were  tlescrihed 
initialK  as  Mcltinia.  and  classified  according  to  shell  eliar- 
acteristics    such    as    sliape    and    sculpture,     llaldeiiian 

'Current  address:  Department  of  /oologx.  Iii\ei1el)rate  l)i\i- 
sioii.  Field  Museum  of  Xatural  Histon,  14(10  .S.  Lake  Sliore 
Dri\e.  Chicago.  IL  60605  US.\:  rmintonta fieldiimseinii.org. 


(1S40I  eri'cteil  Lilliiisin  .ind  dcsignalcd  /,.  ociiii-ulald 
llaldeiiian.  1.S40.  .is  llie  t\p<'  tor  tlie  genus  and  lati'r 
erected  AniHlniiid  (1S41  ).  I'resi'Uce  of  posterior  and  an- 
terior calluses  united  Aiiiiitniud  and  IJlhasia.  The  pri- 
inan  characters  separating  the  genera  were  that  .\/(g(- 
hviiui  shells  were  spinous  and  had  apertures  with  an 
anterior  sinus,  while  l.itliasiti  shells  lacked  senljiture  and 
the  aperture  was  not  as  distinctly  channeled  in  front  a.s 
the  t\pical  Aniiilri'iiKir  iTnon.  1873).  Goodrich  (1921) 
supported  Filshn  and  Rhoad's  ( 1S96)  reduction  ol  A/i- 
'^itrcuui  under  LilluisUi.  and  IJlliiisia  snhseciuentK  has 
stood  as  a  single  genus.  Cloodricli  ( 19401  recognized  lour 
si'parate  groups  wilhin  lAtluisin.  Iiased  priinarlK  on  |)e- 
culiarities  ot  shell  siulpture.  In  tlu"  1970s,  authors  sug- 
gested taxouoniie  revisions  ol  the  genus,  placing  all 
nieiiiliers  in  /<)  (Da\is.  1974)  or  Flfiiroccni  (Stansl)en', 
1971;  Stein.  197S).  Bnreli  and  Tottenham  (19S0)  rec- 
ognized Lithasia  scnsii  stiirtc  and  Aiiiiitrciita  as  subgen- 
era of  Lithasia  based  on  position  ol  sculpture  on  the 
body  whorl  (Burcli,  19S2),  and  not  according  to  the  orig- 
inal diagnosis,  while  Tnrgeon  et  al.  (1998)  followed 
Hurcli  in  recognizing  iJlhasia  as  one  gemis. 

.Vutliors  lia\"e  liistoricalK  assembled  pleurocerid  geii- 
ir.i  based  on  grouping  taxa  with  shared  shell  characters. 
Such  is  the  pri'Nalent  ajijiroach  found  in  the  literature 
published  oxer  a  |)eriod  ol  150  \ears,  and  no  aualxses  ot 
these  characters  exists  lor  IJlhasia  or  an\  other  pleu- 
rocerid genus.  This  max  be  a  result  ol  the  lack  ot  nni- 
lorni  iiilormation  louiid  in  llie  literature.  Diflereiit  aii- 
lliiiis  rarcK  proxided  eomjiarable  levels  ol  (|nalitati\(' or 
(|iuuititatixe  data  in  their  original  descriptions,  and  sel- 
dom used  terms  and  expressions  that  max  or  max  not 
liaxe  the  same  ilescriptixe  counotations  (e.g..  tapering 
xersus  broadly  conic  shells),  making  it  dilficiilt  lor  read- 
ers to  draxx  comparisons  betxxei'U  xxorks.  Manx  descrip- 
tions were  based  on  one  or  a  lex\'  shells,  jiixeniles,  or 
partial  shells  gixeii  to  the  author,  and  radulae  xxere  not 
included  in  these  descriptions.  FinalK',  descri|)tions  were 
subjectixe  based  ou  the  experience  ol  the  author  the 
amoiiut  ol  xariation  tliex  accepted,  ami  their  imderstand- 
lug  ol  the  other  taxa  in  the  literature.  (Jixeii  the  current 
state  ot  Ireshxxati'r  mollusk  tayoiiomx  in  general  and 
pleurocerids  specihcalK,  and  the  fact  that  moiphological 
characters  are  still  usi'd  to  coufinn  taxon  identity,  anal- 


?:v'e  40 


THE  NAUTILUS.  \ol.  116.  NO.  2 


Figures  1-8.  Illiistration.s  ol '.selected  iharaeters  anil  charaetei  slates  iisid  m  the  eladistic  aiuiKsis  niljflmsin.  1.  Lithasia  annigcra 
sluming  lusilonn  aperture  (2:2),  and  presence  oT  aiiteriiir  (7:1  '  and  posteiKii  i(i:l  '  calluses  (in  the  cehnnelhi.  2.  Litliasiii  vcrniri'sii 
.showing  nodulose  brxK'  whorl  (5:1 )  and  poslenor  lenij;llienni<;  el  ;ipeitiiic  larmw:  III:  I  '  .'?.  Alisenee  el  cusp  ne\t  In  lalenil  tooth 
(13:0).  4.  I'lcseiiee  ol  cusp  next  to  lateral  tooth  iarrow:  l.'5:li.  5.  Lamellar  main  lateral  cusp  i  19:11'.  6.  Kectan^iilar  iii:ini  Literal 
cusp  (19:1  I.  7.  Trapezoidal  main  lateral  cusp  (19:2).  8.  Trian<;nlar  main  lateral  cusp  (19:3). 


y.ses  ol  tliese  cliaraet<'rs  cuiploNiny;  modern  tccliniqiies 
is  pnulciil.  Moileni  methods  orpiivlogeuetic  ssstematies 
ha\e  shown  that  some  moqiliological  trails  do  not  sup- 
port historicalK  aeeepled  iiiolhisk  ciassifiealioiis  KJral, 
2()()();  Lydeardet  al..  2000).  11  analv.ses  of  moipho!o<j;ieal 
characters  do  sup])ort  emreiif  tuNonomie  h\|K>llieses. 
tluMi  the  classification  oi  these  groups  can  he  sta!)i!i/ed. 
If  not.  our  approacli  to  iisinsi  diaiinostic  cliaractcis  lor 
those  groups  unist  change,  j'lnlogenetic  taxonomies  of 


this  kind  Iiunc  iieen  advocated  in  several  studies  (e.g.. 
de  (,)uein)/  and  Caulhier.  1090.  1092,  1994;  Biyant, 
1996;  .Sereno,   1999;  l,\de;n-d  et  ;il..  20001. 

l''urther  complic;iting  this  |irol)leni  is  th;it  pleiirocerids. 
Ill  a  simihu'  hishioii  to  wIkiI  h.ippens  to  nuun  other  Iresh- 
watei  inxertehratc  groups,  ;ue  experiencing  (k'clines  in 
llieii  iiumher  ol  species  iuid  iiidividuiils  eiuised  In  river 
impiiimilmenl.  haliit;il  di'gi";id,iti(iii,  and  poor  laiid-nse 
practices  I  Stein.   i9T(i;   Hogiiii  et  al,   1995;   l,V(k';u-d  et  ;l].. 


R.  L.  Minton,  2002 


IVc  41 


1997).  For  example,  in  tlie  Moliile  Basin,  one  ticiiiis  (Cw/- 
roto})ia)  iuul  approximately  31  other  species  an'  prcsnniei! 
extinct  (Stein,  1976;  Bogan  et  al.,  1995;  L\(leanl  and  Max- 
den,  1995;  Lydeard  et  al.,  1997).  Even  with  the  loss  ol' 
di\'ersit\'  in  the  tamiK',  onK'  five  of  156  ri'cogni/.ed  pk-n- 
rocerid  species  (Turgeon  et  al.,  199.S)  are  listed  as  either 
end;mgered  or  threatened  as  of  Ma\.  2001  (U.S  Msh  and 
Wildlife  Senice).  It  nioiphologN-  ;ilone  is  to  he  used  in 
iilentifieation  ol  these  imperiled  taxa,  then  anaKses  ol  these 
characteis  gain  e\en  more  importance.  .\  lack  of  sncli  anal- 
yses can  hinder  efforts  to  recognize,  manage,  and  consciAe 
distinct  taxa  (\\aples,  1991;  Mawlen  and  Wood.  1995) 
within  these  affected  groups. 

Tile  goal  ol  this  stndx  is  to  compile  shell  and  radula 
characters  from  Lilluishi.  anaKze  them  using  cladistic 
techniques  in  ordei'  to  test  historical  and  modern  clas- 
sifications of  the  genus  antl  its  species  composition,  aiul 
to  determine  possible  relationships  of  the  genus  ;uul  its 
taxa  to  other  pleurocerids  based  on  those  anaKses.  Po- 
tential changes  to  the  taxonom\-  of  Lifh/isin  based  on 
these  analyses  and  the  utilit\'  of  using  these  characters 
in  pleurocerid  classification  are  discussed. 


MATERIALS  AND  METHODS 

Specimens  for  die  study  were  either  collected  li\e  oi'  bor- 
rowed from  mu.seum  collections  (Appendix  1 ).  Slu'll  ch;u- 
acters  were  taken  direcd\  from  specimens.  Radnlae  wt-re 
extracted,  cleaned,  and  prepared  according  to  the  method 
described  bv  Holznagel  (1998),  viewed  using  a  ilitaclii  S- 
2500  .scanning  electron  microscope,  pliotographed,  and  an- 
iilvzed.  For  Liflw.siii.  at  li-ast  one  represeiitatix  e  from  each 
recognized  species  and  subspecies  (Bnrcli  and  'lollcnliani. 
19S0)  was  includeil.  Spetiuiens  of  selected  taxa  iipii'scnt- 
ing  five  odier  extant  pleuidcerid  genera  (EUuiul  /c  /"g". 
Leptnxi.s,  Plcuroccm)  were  al.so  inclntled  (.'\ppendix  1 ). 
Data  consisted  of  a  matrix  of  25  cliaracters  (.\ppendix  2, 
Figures  1-S)  coded  as  either  binan  or  multi-state  (Appen- 
dix 3),  and  an;il\y,ed  plnlogeneticalK  under  maximum  par- 
.sinionvwitli  NONA  2.0  (C;olob()ff',  1998)  n.sing  the  follow- 
ing .settings:  nnonk'red  data,  100  replic;ites,  with  /;/gf;  s'd- 
iciila  and  Mrlaiuiidcs  luhcrculald  Miillen  1774.  as  out- 
groups.  //(g(;  is  basal  to  the  rest  ol  the  Xortli  ,\nieriean 
Plenroceridae  (Holznagel  and  L\'deard,  2000)  and  M.  Iiih- 
crcnlata  was  chosen  as  a  more  distant  outgrou|).  jackknile 
analysis  (37%  deletion,  1000  iterations  of  10  i-eplicates 
each)  was  performed  in  XAC  (Fanis,  unpublished:  F;uris 
et  al.,  1996)  to  test  the  stabilitx-  of  the  data  A  strict  con- 
sensus tree  mapped  with  characters  was  prodnc-ed  with 
Winclada  0.9.99m24  (Xixou.  1999).  The  anaKsis  was  run 
twice,  once  using  shell  cliaracters  alone  and  once  with  iill 
ch;u-acters  combineil.  Because  most  pre\ious  classifiealions 
(e.g.,  Tnon,  1873;  Cioodiicli,  1940)  were  based  on  sliell 
characters  onK;  tlie\'  were  anaK'zed  sepaniteK  and  com- 
bined witli  radnki  data. 

Once  the  classification  li\potliesis  was  establislied.  it 
was  compared  to  fixe  diflereut  classification  selicnies 
proposed  b\-  prexious  authors  (Appendix  4': 


:\.  bitliasia  lepresenls  a  single  genus.  This  assumption 
lolioxxs  the  cuiR'ut  iTurgeon  et  ;il..  1998)  xiexxofthe 
genus. 

B.  Ijtiuisia  represents  a  single  genus  comprised  of  txxo 
subgenera,  Lilliiisia  scif-.ii  strirlo  anil  Aw^itrruui. 
sciisii  Hiurh  d\n\  Tottenham  il980).  This  classifica- 
tion is  conimoiilx  used  as  ;i  st:uting  point  in  pleu- 
rocerid studies  leg.,  i.xdcard  et  al,.  1997:  Holznagel 
and  Lxxk'iu-d,  2000).  Bnrch  and  Tottenham's  (I980» 
genera  and  subgenera  differ  in  species  composition 
Ironi  those  ol  TiAon  ( 1873). 

O.  Lithasia  repicsents  a  single  genus  comprised  of  four 
species  groups  based  on  peculiarities  of  the  nodulous 
sculpture  ((ioodrich.  1940).  Lithasia  luihriclili 
()lench.  1965.  had  xet  to  be  described  and  is  includ- 
ed in  (irou|i  3  based  on  (dench  i  1965'  allxing  it  to 
Lithasia  ti'micasa  i  jiafiuesque,  1820'. 

D.  Taxa  presentlx  included  in  Lithasia  bi'long  to  one  of 
tluei'  genera:  Lithasia,  Ani^itrciiia.  or  Annihisa.  scii- 
sii Trxon  (1873).  In  this  case,  the  original  descriji- 
tions  <il  Lithasia  ;ind  .\ii<iitrciiia  ;n'e  used  to  group 
taxa  based  on  shell  characters.  Tnon  failed  to  in- 
clude Lifliasia  ciiiia  (Lea,  1868),  and  Lithasia  hii- 
hiichti  luid  xet  to  bi-  described.  Both  species  are  in- 
cluded in  Ti-xons  Aiiiiilrciiui  based  on  their  nodulous 
shell  sculplure.  Lithasia  i^ciiiciilata  jiiirj,iiis  i  Lea. 
1852)  =  Aiiciilosa  piu^uis. 

E.  Taxa  presentlx-  included  in  Lithasia  should  be  con- 
sidei'ed  speiies  ol  lo  based  on  dexelopmeutal  char- 
acters, SCIISII  Daxis  (1974'.  lo  Lea,  18.31,  has  prece- 
dence oxer  Lithasia  Haldeman,  1840,  uml  Aufiilrcma 
Haldeman,  1841. 

.\  sixth  scheme.  Steins  (1978b'  recommendation  that  all 
Lithasia  be  considered  Piciiroccra.  is  not  treated  here  as 
it  IS  nomiiicl.iliiial.  not  taxoiioinic.  and  has  since  been  re- 
solxed  b\  the  liilernational  (>)iiiiiiission  on  Zoological  No- 
menclature's decision  to  make  Flciiroccra  anitiis  tlie  tx]^' 
species  ol  the  genus  i  Melxille.  1981:  see  discussion  in  Bo- 
gan ;ind  I'armalee.  1983i.  (Characters  supporting  relation- 
ships in  the  paisiinoux  analxsis  xxcre  com|)ared  to  charac- 
ters that  groiijied  spei  ies  in  the  other  ckLssific'atioiis. 

RESULTS 

Maximuni  p;irsiiiioiix  analxsis  of  shell  cliaracters  alone 
xielded  372  trees  of  27  steps  (Figure  9).  Lithasia  was 
rendered  non-inonophxktic  b\  the  placcMiient  of  Lith- 
asia 'Uiiiiiilata  jiiii'^iiis  in  the  clacie  ol  Lcploxis  species 
;iiid  the  piac-emeiit  ol  Lithasia  ohoiata  (Sa\\  1829)  in  a 
polxtomx of  I /(I  )  some  Lhiiiia  +  {pin<iiiis  -^  l^cptoxis) 
+  remaining  Lithasia'.  .\nalxsis  ol  all  characters  xielded 
20  trees  ol  107  steps  that  rendered  Lithasia  non-mono- 
plixlclic  i  Figure  10'.  Lithasia  <iciiiciihita  ])iiiiiiiis  speci- 
iiieiis  were  basal  to  a  clade  ol  Lcptoxis  s|)ecies  supported 
bx  a  te:n'drop-slKiped  aperture,  and  Lithasia  oboiata  xxas 
iiesled  belwecn  ckules  cA  Ph'iiroccra  and  LUiiiia  species 
near  llie  base  ol  the  tree.  The  remaining  Lithasia  taxa 
loiined  a  clade  xxitli   In  supported  bx   three  cliaracters: 


Paw  42 


THE  XAUTILl'S.  \oI,  IKS.  Xo,  2 


Melanoides  tuberculata 
— —    E.  caelatura 
— ^    E.  hydei 
^^   Juga  silicula 


81 


1 


P.  canaliculatum 
P.  prasinatum 
P.  walkeri 


Lithasia  obovata 
^^    lo  fluvialis 
E.  olivula 
E.  alabamensis 


65 


Lithasia  geniculata  pinguis  Duck 
Lithasia  geniculata  pinguis  Collins 


Leptoxis  ampla 

-  Leptoxis  crassa  anthonyi 

-  Leptoxis  plicata 
Leptoxis  praerosa 
Leptoxis  taeniata 
Leptoxis  virgata 

— — -    Lithasia  armigera  Ohio 
— i^^—    Lithasia  armigera  Stones 
^^^—    Lithasia  armigera  Harpeth 
-^^^    Lithasia  armigera  Wabash 
^^^—    Lithasia  duttoniana  Duck 
^^    Lithasia  duttoniana  Harpeth 


69 


Lithasia  geniculata  fuliginosa  Buffalo  1 
Lithasia  geniculata  fuliginosa  Buffalo  2 
Lithasia  geniculata  fuliginosa  Harpeth 
Lithasia  geniculata  fuliginosa  Duck 
Lithasia  geniculata  fuliginosa  Red 
^^    Lithasia  jayana 
Lithasia  salebrosa  florentiana 
Lithasia  salebrosa  subglobosa 

Lithasia  geniculata  geniculata 

-  Lithasia  hubrichti 

-  Lithasia  salebrosa  salebrosa 

-  Lithasia  curta 
^—    Lithasia  lima  Bear  Creek 
^—   Lithasia  lima  Elk 


T^ 


Lithasia  verrucosa  Ohio 
Lithasia  verrucosa  TN 
Lithasia  verrucosa  Wabash 
Lithasia  verrucosa  White 


Figure  9.     Chuiistic-  iiiuilysis  ol'  Lithasia.  Strict  consensus  of  tiie  372  most  parsimonious  troi's  (27  steps:  CI  =  0.74)  ijencratecl 
usiTii;  siiell  clianic-ters  aloia-.  Jackkiiile  \alnes  >  63%  aliove  nodes,  number  of  imamhisiuous  synapomoipiiios  below  nodes. 


Mintnn.  2(102 


Patie  43 


^-  Melanoides  tuberculata 
—  Juga  silicula 

81  T~~  P.  canaliculatum 

2  I      I P-  prasinatum 

* —   P.  walkeri 
t—  Lithasia  obovata 

^^  E.  olivula 


77 


E.  alabamensis 
E.  caelatura 
E.  hydei 
—  Lithasia  geniculata  pinguis  Duck 

Lithasia  geniculata  pinguis  Collins 
Leptoxis  plicata 

Leptoxis  virgata 

Leptoxis  crassa  anthonyi 
Leptoxis  praerosa 
Leptoxis  ampla 
Leptoxis  taeniata 
—  Lithasia  armigera  Ohio 
Lithasia  armigera  Stones 
Lithasia  armigera  Wabash 
Lithasia  hubrichti 

Lithasia  armigera  Harpeth 

Lithasia  duttoniana  Harpeth 

-  Lithasia  curta 

-  Lithasia  lima  Elk 

-  Lithasia  lima  Bear  Creek 
Lithasia  verrucosa  Ohio 

-  Lithasia  verrucosa  TN 

-  Lithasia  verrucosa  Wabash 

-  Lithasia  verrucosa  White 

—  Lithasia  geniculata  fuliginosa  Buffalo  1 

Lithasia  geniculata  fuliginosa  Harpeth 

—  Lithasia  salebrosa  florentiana 

—  Lithasia  geniculata  fuliginosa  Duck 

—  Lithasia  salebrosa  subglobosa 
—  Lithasia  duttoniana  Duck 

—  Lithasia  geniculata  fuliginosa  Buffalo  2 

Lithasia  salebrosa  salebrosa 

I —  Lithasia  geniculata  fuliginosa  Red 
^H        r^  Lithasia  geniculata  geniculata 

I        I Lithasia  jayana 

' lo  fluvialis 


Figure  10.     (,'lailistic  an.il\sis  of  Lilhasia   Strict  consciiMis  of  llu-  20  most  p.irsiiiionioiis  trees  i  lOT  steps.  CI  =  ()..374)  <;eneratecl  using 
shell  aiul  radula  cliaraeters  conililiied.  j.iekknite  \alues  =r  (i:5'<  alune  iKides.  miiiil)er  "I  uiiaiMhi'.ni()iis  .sviiajToiiioiphies  below  node.s. 


Page  44 


THE  NAUTILUS.  \\>\.  Ilfi.  No.  2 


tusifbnii  aperture  (c-liaract(>r  2:  state  2),  posterior  i-allus 
on  aperture  l6:l).  and  slij^ht  anterior  eanali/.ation  ol  the 
aperture  (S:l).  Despite  being  well  resolved,  little  snppoit 
for  an\'  elade  existed  as  evideneed  1)\  low  jaekknili'  val- 
ues. Three  other  plein"oeei"id  genera.  Eliiiiiti.  Ij'jitoxis. 
and  Vliiiroccru.  \w\v  all  I'eeovfri'd  as  ni(iiiiipli\  letic. 

DISCU.SSlOX 

The  c'ln'rent  taxonomv  ol  pleuroeerids  is  based  on  shell 
cliaracteristies.  and  most  work  on  the  faniilv  has  foeuseil 
on  these  characters.  An  extensive  literatnn'  exists  lor  the 
laniilv,  with  most  works  being  either  wholK  tlescriptive 
or  taxonomic  shullling  taxa  among  groups.  This  studv 
stantls  as  the  first  cladistic  treatment  ot  all  cnrrentiv  rec- 
ogni/eil  l.itliiisiti  species  and  their  relationships  to  other 
pleuroeerids  based  on  shell  and  radula  characters. 

Neither  anaKsis  completely  recovered  the  five  classi- 
fications being  comparetl.  In  both  [ilivlogenetic  treat- 
ments Lilha.sia  taxa  did  not  tnrm  a  single  group,  which 
refutes  the  taxonomies  of  Bnrcli  and  Tottenham  (f9S()), 
Goodrich  (1940).  and  Tnrgeon  et  al.  (199S).  Tiyou's 
(ISTo)  groupings  ol  the  cnrrentiv  recognized  species  of 
Litluisia  was  also  not  supported,  as  sculptured  and 
smooth  taxa  did  not  group  separatelv.  Davis's  contention 
that  LUhasia  species  should  be  considered  as  members 
of  /()  was  partiallv  supported  in  the  parsimonv  aualvsis 
of  all  characters,  where  lo  was  nested  deep  in  a  clatle 
containing  most  Lithtisiii  taxa. 

The  const'nsns  ti'ees  suggest  tliat  sliell  characters  alone 
do  not  recover  cnrrentiv  or  historicallv  recognized  groups. 
However,  shell  and  radula  characters  combined  can  be 
u.sed  to  recover  jilenrocerid  genera,  but  do  not  resolve  spe- 
cies level  identilv  well.  In  ihc  lota!  chaiattei  an;Jv,ses,  EJi- 
m'ui.  lA'j)l(>xis.  and  licuioccra  taxa  all  grouped  in  their  re- 
spective genus.  Onlv  hvo  IJthasid  taxa  grouped  awav  Ironi 
the  others,  suggesting  that  these  two  species  mav  be  mis- 
placed. lAtliaski  >!fnicnlalii  piii'^iiis  was  placed  iu  h-jiloxis 
(  =  Anailosd)  by  Tnon  (1S64I,  and  subse(|uentl\  [ilaced 
in  LilJias'm,  vvliere  it  has  icpreseuted  the  headwaters  lorni 
of  the  ^ciiinildld  '^ctnaddld-fnliiiiiHKd-p'uiiiiiis  complex. 
Based  on  the  moiphological  cliaracters  examined  here.  /, 
i^eniculutd  piiiiiiiLs  should  be  classified  as  a  species  (i\'  Ia'j)- 
toxis.  lAtlui.sid  ohovdid  is  the  onlv  species  in  the  genus  thai 
(X'cnrs  in  the  Creen  River  drainage  of  Kenlnekv  and  has 
included  njanv  nominal  forms  that  are  (jneslionablv  l.illi- 
asia.  L.  ohoiaUi  shells  lack  the  calluses  on  liic  apci-tmc. 
fiisifonri  a])ertures.  and  have  radulae  most  similar  to  Elimiii 
species.  Though  the  plivlogenetie  aualvsis  suggests  alloca- 
tion o(  /..  iilnnald  to  a  separate  genus,  I  belii've  the  ad- 
dition ol  more  EliinUt  and  Fleiirorcrd  species  or  more  mor- 
plioiogicid  chaiaclers  In  ihe  aualvsis  would  icsolve  its  ge- 
neric designation.  The  remaining  Litluisia  species  would 
lie  cousiilered  h>.  as  hi  is  the  oldest  name  for  that  clade. 
As  such,  lo  would  be  dia'gnosed  bv  having  a  fusiform  ap- 
erture (2:2),  posterior  i  alius  on  columella  (6:1),  and  for- 
mation of  canal  of  the  aperture  (S:l-2).  This  change  would 
reflect  the  opinions  ol  fJavis  '  1974)  regarding  the  two  gen- 
era. Within  Litha-sia,  liowevei',  individual  species  weic  not 


recovered,  and  continued  aualvsis  is  re(]uired  to  elucidate 
diagnostic  characters  al  the  species  level.  None  ol  the  five 
current  and  historical  tlassifications  ol  Eitluisid  evaluated 
iu  this  studv  are  completi'lv  cousisti'iit  with  the  aualvsis. 
OuK  the  diagnosis  ol  Bnrc-li  (19S2)  is  partiallv  supported. 
A  posterior  cdlus  on  the  columella  (6:1 )  and  the  fonnation 
of  anterior  canal  of  die  aperture  (S:l-2)  unite  all  Litlwsia 
(minus  ff'nicuUitd  pin^iiis  and  ahot nla  plus  la  i  in  the  corn- 
binetl  character  aualvsis. 

This  studv  offers  evidence  refuting  pri'vious  notions 
that  shell  and  radula  characters  have  limited  utilitv  in 
recognizing  pleinocerid  groups  and  supports  the  use  of 
these  characters  in  defining  pleurocerid  genera,  in  the 
most  inclusive  treatment  of  pleuroeerids  to  date,  Tnon 
(IS73)  offered  an  extensive  discussion  on  tlie  use  and 
validitv  of  shell  characters  iu  separating  t\\v  various  gen- 
eia  and  species  in  the  group.  Tnon  recognized  that  shell 
characters  can  van'  greatlv  and  looked  towards  the  use 
ol  other  anatomical  characters  to  separate  "natural  gen- 
era" and  discover  corroborativ  e  sliell  characters  for  these 
groupings,  (a)oilrich  (1940:  1)  noted  that  shell  charac- 
ters "once.  .  .considered  inunutable  have  proved  to  be 
secondan'  and  more  or  less  evanescent'.  Tnon  (1873: 
liii-lv )  figured  some  ol  Troschel's  illustrations  ol  radulae 
and  conunentetl  on  Stimpson's  obsenatious  that  sliell 
and  ladnla  characters  seem  to  unite  pleuroeerids  but  do 
little  lo  sejiarate  constituent  genera  and  species.  It  is 
triH-  that  gastropod  radulae  (Padilla.  199Si  and  tresh- 
vvatei'  mollusk  shell  characters  can  be  plastic,  olten  ex- 
hibiting clinal  variations  (e.g.,  .Adams,  1900,  1915;  Ort- 
maun,  1920),  ami  therefore  potentiallv  contrilinting  ho- 
moplasv  to  phvlogenetic  aualvses.  Adtting  solt  anatomv 
characters  to  a  studv  such  as  this  might  tlieoreticallv  im- 
piovc  the  resolution  ol  the  analvses,  lint  manv  eharactei's 
lre(|ueutlv  used  to  delineati'  taxa  van  little  among  pleu- 
locerids  (Dazo,  1965)  and  in  phvlogi'iietic  reeonstna- 
tious  shell  chaiacters  often  are  less  homoplastic  than  an- 
aloniic;il  cluirai'ters  i  Schander  and  Sundberg,  2001). 

The  results  given  here  are  consistent  with  studies  em- 
plovnig  molecukn'  methods  (Lvtleard  et  al..  1997:  Lv- 
di';n-d  el  :il.,  I99S;  llolznagel' and  Uvdeard.  2000)  to 
idenlilv  [)Ieuiocerid  genera.  In  tlii'se  works.  Eliiiiid  and 
I'lcuroccni  represent  natural  groups,  and  the  uioqiho- 
iogic:il  characters  siippoi't  both  genera.  However.  Lv- 
(leard  (I  al.  (1997)  and  llolznagel  and  Lydeard  (200()) 
showed  lli:il  Lilhdsid  :md  Eiploxis  are  uon-mono]iliv  let- 
ic, ihoiigh  the  eombmed  iiiorjiliological  analvsis  given 
here  supports  the  reeoguitioii  ol  Ecj)l(>xis  as  a  natural 
giou|).  \  (horough  moleenlai' sindv  ol  IJllidsid.  including 
all  ol  its  nomin:il  species  and  lornis.  \\\\\  pro\  ide  \  ;ilu,il)ie 
inloiiualiou  on  the  species  composition  ol  Lillidsia.  anil 
subse(|n('iil  ehaiacler  :iiialvsis  will  lielp  driiui'  ihe  diag- 
nostic le:ilin'es  ol  tlii'  genus. 

.ACKNow  i,i;i)(:\ii:nts 

I  would  like  to  Ihank  ,S,  Ahlsledl.  |.  ( ;,uner,  and  I),  \eelv 
lor  s|)((iiiirns.  \I.  (ilaubreeht.  I'.  I  lariis.  ('..  lAtk'ard,  and 
v..  Strong  provided  heljilul  comments  on  the  manuscript. 


R.  L.  Miiitnii.  2002 


Paiif  45 


W.  Hol/iuiiifl  ami  |.  NiinlcN'  lussisted  witli  radula  prepara- 
tion and  illustration,  and  \'.  Albert  pnnided  access  and  in- 
stniction  to  NONA,  W'inclada.  and  -V\(".  A  Resi'arcli  (  aant 
from  Concholc joists  of  Anienca  Inndetl  this  project. 

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water  mussels  (Naiades).  Proct'edings  ol  the  .Vmeriean 
Philosophical  Society  59(4):  269-312, 

Padilla,  D.  K.  1998.  Inducible  phenot\pic  plasticity  of  the  rad- 
ula  in  Liiciiim  ((Gastropod;!:  I  ,ittorinid;ie'.  The  X'eliger  2: 
201-201, 

Pilsbiy,  II.  A.  and  S.  N.  Rhoads.  1896.  Conlributions  to  the 
zoologN  ol  Tennessee.  No.  I.  Mollnsks.  Proceedings  of  the 
Academy  of  Natural  Sc-ienees  of  I'hifidelplii.i  IS:  487-506, 

Rafinesque.  C.  S.  1820.  Annals  ol  n:ilnrc,  or  .iniiu;il  s\n<ipsis 
of  new  genera  and  species  of  (Kiniinals.  pfinls.  etc.,  dis- 
covered in  North  America.  Pp.  10    11. 

Redfield.  J.   II.    1854.   Deseriplious  ol   new   species  of  shells. 


.\nnual  Report  of  the  Lvceum  of  N;itural  Histon- of  New 
York  6:  130-132. 

Ree\'e,  L.  A.  1860.  Conehologia  leonica:  or  illustrations  of  the 
shells  of  molluscous  animals.  Lowell  Ree\e,  London. 

Sa\,  T.  1821.  Descriptions  of  the  univaKe  shells  of  the  United 
States.  lournal  of  the  ,\cadem\-  of  Natural  Sciences  of 
Philadelphia  2(1 1:149-179. 

Sa\",  T  1825.  Descriptions  of  some  new  species  of  fresh  water 
and  land  shells  of  the  United  States,  [oumal  of  the  .Acad- 
emy of  Natural  Sciences  of  Philadelphia  5(3-4):  119-131. 

Sa\,  T  1829.  Descriptions  of  new  terrestrial  and  flmiatile  shells 
of  North  America  (continued).  New  Harinon\-  Dissemi- 
nator 2(19):  291-293. 

Schander,  C.  and  P.  Sundberg.  2001.  Useful  characters  in  gas- 
tropod phvlogenv:  soft  information  or  hard  facts.  Svstem- 
atic  Biolog>- 50:  136-141. 

Sereno,  P.  C.  1999.  Definitions  in  ph\logenetic  t;L\onom\:  cri- 
tique and  rationale.  S\steniatie  Biologx  48:  329-351. 

Stansben;  D.  H.  1971.  Rare  and  endangered  freshwater  mol- 
lusks  in  Eastern  United  States.  In:  Jorgenson,  S.  E.  and 
R.  E.  Shaip  (eds.)  Proceedings  of  a  ,s\mposium  on  rare 
and  endangered  mollnsks  (naiads)  of  the  U.S.  U.S.  Fish 
and  Wildlife  Sei-vice,  Department  of  the  Interior,  Wash- 
ington, pp.  5-18f.  50  figs. 

Stein,  C.  B.  1976.  Gastropods.  In:  Endangered  and  threatened 
species  of  .\labaiua.  Bulletin  of  the  Uni\ersit\  of  .Alabama 
Museum  of  Natural  HistoiT  No.  2.  Tuscaloosa,  pp.  21—11. 

Stein.  (.'..  B.  1978b.  (xiuunents  on  the  proposed  designation  of 
a  t\pe  species  lor  rlciiroccni  Rafinesque.  1818.  Bulletin 
ol  Zoological  Nomenclature  34:  196-197. 

TiAdii-  ( '.  W-  1864.  S\iion\ni\  of  the  species  ol  Strepomatidae, 
a  lannK  of  fluvatile  \Iollusca.  inliabiting  North  .America. 
Proceedings  of  the  .\ca(kMn\  ol  N:ifunil  Sciences  of  Piiil- 
adelphia  16:  24-48,  92-104: 

TiTon,  G.  W.  1873.  Land  and  Freshwater  Shells  ol  North 
America.  Part  I\'.  Strepomatidae.  Smithsonian  .Miscella- 
neous t^ollections  16(253),  pp  i-K   +  1—135. 

Turgeon.  D.  D..  [.  F.  Quinn  |r.,  .X.  E.  Bogan,  E,  \'.  Coan,  F. 
G.  Ilochberg.  W.  G.  L\ons.  R  M.  .Mikkelsen,  R.  J.  Neves, 
C.  F  E.  Roper,  G.  Rosenberg.  B.  Roth.  A  Schletenia,  V. 
G.  Thompson.  M.  X'ecchione  and  G.  D.  Williams.  1998. 
C!ornmon  and  scientific  names  ol  aquatic  invertebrates 
from  the  Lhiited  States  :uid  C'anada:  Mollnsks  (2nd  edi- 
tion), ;\meriean  I'lsheries  Societx  Special  I'ublication  26, 
52(i  pp.  r,  S,  Fish  and  Wildlile  Senice  Threatened  and 
End:ingered  Species  S\steni,  littp:  (•n(l,in'4ered,lws,<rov. 
Queried  M:i\   S.  2001, 

Wapli's,  I'l  S  1991,  I'.ieilie  s:iliiiiiu.  Oik  orluimlius  spp,.  :in(l 
the  (Icliiiitidn  ol  species'  under  the  lMidan'.;<i"ed  Species 
\(l,  M.iinic  l-'i-slieries  Re\iew  53:   II    22, 


R.  L.  Miiitoii.  2002 


raui'  47 


Appendix  1.  Ssstciiiatic  list  dl  la\a  used  iii  iIk   i  I.kIisIic  analxsis  nl  tin-  minis  LUliasia  i  ii  =  I  lor  rarli  t.iMiii '.  ( lissification  follows 

Turgeoii  v\.  al.  (UWS):  la\a  aic  iiaiiicd  as  m  Hiinli  (liJSOi,  C pldc  ImalitN  iiitoniiatioii  is  axailalilc  Ikhii  iIic  aiillior,  I'MNII  — 

Field   Miisi-iiiii  nl    Xatmal    llislon     I\1IS    -llliiiiiis   Natural    ilistorx    Siirvi'V:   NCSM  — Nmlli   Carnliiia  Stair   Miisiiiin  iil   Xaliiral 
Sciences;  UAti — Lluivcrsitx  ol  Alaliaiiia  Castropinl  ( .'ollectioii;  I'MM/ — l)iii\cisil\  of  \licliii;aii  Miisciiiii  nl  /iiolii'j;\ 


Taviii 


Localit\ 


( A)llrctiiiii 
niinilxM" 


FaiiiiK   Plriirocrridae 
(aMiiis  Eliinid 

F.    (llilhdliuil.sis  (Lea,   ISdll)) 

E.  ciicliitnrd  iinliiliini  iKnAe.  ISIidi 

K   hi/ilci  (Cloiirad.  IS:54lii 

K.  (tliiiilii  (Cimrad,  IS34a) 

Cleniis  Id 

I,>  fliiiialis  (Sa\.  1S25) 

Ca-iiiis  jiii^ii 
I    silinilii  iC.nild.   1S47) 

Cieniis  Lfj)loxi\ 

L.  init])l(i  (AntlmiiN.  1.S55) 

L.  cnissii  (iiitlioin/i  ( l^edfield,  IS54) 

/.,  /)//(■(//(/  (Conrad.  l.S34li) 

/.  pnicrosii  (Sa\,  IS2I  i 

/,.  Iiiiiiiiitd  ((Jinratl.  1S341)) 

/„  tinj^atd  (Lea.  lS41al 

(ieniis  Lilluisiii 
L.  iirini''!  Ill  I  Sa\.   1S21 ) 


/..  aiiia  (Leu.  1868) 

L.  iliittimiiniii  (Lea.  1841a) 

I,    •j^i'iiiniliilii  ■j^iniiiiliilii  {\\Mv\iidt\.  1S4I) 
E   >^ciiiciiliitii  fiili^iiKKii  (Lea.  18421 


/., 


IciiifuJiitii  jiiiipii'i 


(Lea,  1852) 


E-  hnhiirhti  CIcikIi,   1^65 
E.  jaijdHil  (Lea,   184  llil 
E   liiiui  I  (  'niirail,  18(34a) 

E   oh, It, ilii  iSa\.  182V)t 
E  MilcliiiKii  sidchi'iKii  (Conrad.  1834a) 
L.  salrhnisii  fh>nuli/iiui  (Lea.  1841a) 
L.  salfhntsa  siih^Lihosa  (Lea.  LSfila) 
L.  verrucosa  (Hafiuesque,  1820) 


Genus  Plrurocrii 
P  rinudiiidaiuiii  filuin  (Lea.  1845) 
/'  pnisiiiiiliiiii  i(!oiir.id,  lS34a) 
/'  uidki'ii  C.H.drieli.  UJ28 

FauiiK  Tlilarldae 
(  a'lllls  M, 1,11101, lis 

SI   liihintiliilii  i.Mullcr.  17741 


( !uosa  l\i\er,  ('oosa  ( 'o..  M  , 
(111  Kii  linen  1  (   reck,  (  'all  HI  11 II  (  (i,,  .\L 
Black  Warrior  Hiver.  Jericisi.ii  Co,,  AL 
Alabama  Ri\er.  .Monroe  (!o,.  .\L 

llolstoii  Rl\ir   Siilli\aii  Co     T\ 

Oak  Creek.  Hciitmi  (  o    OK 

Liltic  Cahalu  Uimi,   Kilili  (  ii      \L 
Sci|natcliie  Ki\er.  Marion  ( !o,    AL 
Black  Warrior  Hi\cr,  |ellcrsoii  (  o  ,   \L 
llaipetli  Hi\ri,  OaMclsoii  (  ji    '!'\ 
(  lioccnioceo  (deck,    I'alladi'^a  ( (o.,  .\L 
(diiicli   l!i\ei,  I  laiirock  ( 'o,.    IN 

Wal.asli   l;i\ci    Wliilr  Co     11, 

Oliio  Hixcr.  Massac  Co,,   IL 

h'.asi  I'oik  SloiHs  i;i\(i    Uiitlicrlord  Co    TX 

llarpitli  ItiMi    (  :lii-,itliaiii  (  n      lA 

lennessee  l\i\ci.  Lauderdale  (ai..  .\1, 

Duck  Hi\er.  Maun  Co..  T\ 

llarpctli  Ki\er,  Davidson  ('o,,  T\ 

Caiicx   Fork.  Dekalli  Co,.  TN 

Biillalo  Hi\er.  Iluiiiplire\s  Co.,  4"N  (1) 

Red  River.  Rolicrtsou  Co,.   I'N 

Duck  River.  Maun  (;<).,  TN 

RulTaki  River.  Pern  Co,.  TN  (2) 

llaipeth  River.  Davidson  ('o.,  T.N' 

Oillins  River,  Warren  Co.,   TN 

Duck  River.  Coffee  Co,.   1\ 

Ht'i  Black  River.  Hinds  Co.,  MS 

Duck  River,  lliiinplircvs  C.n..  TN 

I'dk  Hiver.  Limestone  ( 'o      M. 

Bear  Creek.  Collierl  Co,,  ,\l. 

Careen  River,  Mel  .r.in  (  n     K\ 

Tennessee  River.  Lauderdale  Co..  .\L 

Tennessee  River.  Hardin  Co..  TN 

Tennessee  River,  ILirdiii  Co,,  TN 

Waliasli  River.  Wliile  (  "     il. 

Olio  River,  Massac  Co,,  II. 

Tennessee  River,  Hardin  ('o,,   IN 

Tennessee  River,  L.iiiderd.ile  (  (o  ,    \l. 

Duck  Ruei    M.iiiiA  Co      IN 
Vellovvleal Creek.  Slielliv   Co,.  AL 
Slioal  Creek,  Lauderdale  Co,.  AL 


Draina'^e  ditcli.  Cainesville.  ,\l,icliua  Co,,  |-'L 


N(:SM-R-4658 
NCSM-l'-465y 
NCS\LI'-46(i3 
NCS.M-l'-4(i(i4 


NCSM 


IfifiT 


NCSM-l'-4(i7() 

NCSM  1'  4671 
NCSM-l'-4(i72 
NCSM-l'-4fi74 
NCS\LR-4(i75 
NCSNM'-4fi7fi 
NCSNM'-4(S77 

INI  IS  2.3628 
INHS  236.32 
UA(;  397 
UAC  572 
UMMZ  2122()() 

vm:  402 

UAC  405 
U. MM  7.51363 
VAC.  406 
UAC;  398 
UAC  403 
UAC  395 
UMMZ  53233 
UAC  407 
UAC;  392 
FMNH  137751 
l^AC  573 
UAC  571 
UAC  570 
FMNH  46219 
UAC;  565 
UAC;  425 
UA(;  416 
INHS  23629 
INHS  23631 
UA(;  427 
LAC  568 

NCSM-I'-46S6 
NCSM-I'-4689 
NCSM-l'-4692 


N(:SM-P-46S2 


ra<re  4S 


THE  NAUTILI'S.  \(.l.  116.  N< 


Appendix  2.   M„n,l,..lo,ic.l  elunacto  a,ul  cluuactc,-  states  used  in  tlu-  clad.stic  anaKs.s  of  the  genus  LUl... 


1    Shell  shape,  l"!  t;li>bose.  1 1 »  conic,  (2)  ovatel)  c v 

0    Aperture  shape. \())  teardrop,  i  1 1  oNate,  Cl)  lusih>nn. 

^'   Sculpture  on  posterior  hodv  whorl.  lO)  none,  (1)  carinate,  (2)  tu hercles. 

t  f  n,e!lialK  on  hodv  whorl.  ,0)  none.  U)  sha.p  angle  on  InxK  f'^^^^^^^^;^^ 

5  Sculptu'v  on  enfre  l,o,h  uhorl.  (01  n,>ue,  (1)  even  lateral  rows  «i  snudi  nodules,  (2)  costate.      ^ 

6  Posterior  callus  ..n  cohunc-lla.  (()>  absent.  (1)  present. 
7.  Anterior  callus  on  cohnnella.  lO)  absent.  (1)  present 

S    Lencnh  of  anterior  eanal  of  aperture.  (0)  none,  (1)  slight,  (2)  elongate. 
9    TwisHna  of  aperture  anteriorly.  (0)  absent.  (1)  present. 
10    Lencnheuingof  aperture  posteriorly  along  bodv  whorl.  (0)  absent.  (1)  present. 

;:■   Sa::f 'i;:;i:^  S: ti  tS^l ^-^^^^o..-..^  ^H^H  l'->.tb,  ^.)  ...ore  than  one-han  shell  length, 
1.3    Cusp  next  to  lateral  tooth  exteriorly.  (0)  absent,  (1)  present. 
14    Shane  of  upper  rachidian  margin.  (0)  convex,  (1)  straight. 

5  nhAvic  th  raHo  of  rachidian.  (0)  tooth  as  long  as  uide   (1)  tooth   ongerhan  v.  le 

,6    Lenath/width  ratio  of  central  rachidian  denticle.  (0)   eiigth  and  uukh  equal,  (1)  longu  than  u.ck. 

1;:  ?±:;:  ^f^;^! .^tl? oifrlnttl^lC^^  -  u.edia,  quarten  (1)  edge  restncted  to  medial  h..  >2,  edge 

s-  EHSU'io '- - 1:^^^^^^^^^       -  =i->-  - '•  -■ '---  "'■  --- 

01  Sgt£:r;:;io:n;:::nS  C5.-fe.i.  ^^.." ;  '-^  --!■  <;  >  ^-f ; ---■  ^"^ "'"■ ' "  ^^^'"  "■"■'' '""  '"*' 

22:  Shaix"  of  leading  edge  of  mmn  lateral  cusp.  (0)  pointed.  ( 1 )  rounded,  (2)  stiaight, 

0.3    Shapeof  marginal  teedi.  (0)  pointed,  (1)  round. 

04    \uniber  of  inner  niar0nal  teeth.  (0)  1^,  (1 )  5-.S,  (2)  more  than  S. 

25'.  Number  of  outer  marginal  teeth.  (0)  1^,  1 1  )  5-S,  (2)  more  than  S. 


Appendix  3.   CMadistic  anal 
ta\(iii  names  where  needeil 


\sis 


IJtlu.sia    Data  matrix  of  t^ixa  and  character  states.  Uixm'  names  loll.m  approi 


iriate 


~  ~  -202010100011011002101112 

I.ilhasia  armfj,,,;,  OInu  2201010100011011012102112 

Lilhasia  armi'^rra  Stones  2202  010100011011001012101 

IMluisw  anm<icra  Haipe  li  2202010100011011001102112 


iJllmsUi  (inniiH-ra  W'abas 


2200111101011011010112112 


Lilhasui  nuiri  2201010100011010020212112 

Lilhasu,  dnllouuma   Dn.'k  2200010100011011013020102 

/.,•/ m.sm  </„»..MJ.,m  1  aipetl,  2200011100011011020211112 

IMhasm  ^■n,rnlala  juhiimos,,   Kullalo  I  2200011100011011021212112 

2200011100011010021212112 
2200011100010011021212112 
2200011100011010120211112 
2220011100011010110111112 


alo 


Lilhasia  iif'nindata  fnliainosa  Bullal 
Lithasin  oniiniliilti  fiili<I,iiu>sa  Duck 
IMluisia  \Sfuinilat(i  ftdiilinosd  llarpetl 
Ijlhasiii  iLcuinilala  fidifliniKd  Hi-d 


lilhasia  i.-nindala  grmn,/«J.  0000000000001011011102112 

.„W,  ^.nndala  ,nu^u.  Duck  0000000000011010111202112 

/,i//w.vm  -cmr„/r;^/  ,.mt;,».s  Colhus  ooonnn  nnnm  1  Oil  0111121 1 2 


IJlliasia  huhiichli 


2220011000011011011112112 
22  02010100011110011111111 


lUlmsu,  jaynnn  2200110101011011011112112 

Ullwsia  hnw    iear  (a.'ck  2200110101011011011112112 

Ulhasm  h,„a   I- Ik  2100000000011011001021112 

.,(  m.s,«  <.to/  0220011100011010020211112 

I,lha.sw  saleh.u,  salrhnsn  2200011100011011021212112 

.,...,«  salehn   ''  tlon.'ann  2200011100011010121212112 

./  ,«.v,«  .v«/.Ws«  .,/,,/,./..«  2200111101011011011112112 

'"""'"  '^'■'"'•'"•'"■"  -V^.'"  2200111101011011011212112 

Ulhasu,  .crn.rosa   IN  2200111101011010011212112 

lilhasu.  vrrnuosa      abash  2200111101011011001102112 

./.;.v,.a.rn,o..s.«Uh>.e  2200000200011010110100111 

/()  jiuiiali'i . 


R.  L.  Mintoii,  2002 


Pam-  49 


Appondiv  3.   (  cmti 


.1 


Li'ptoxix  1 1  mill  1 1 

Lcptoxis  cyiisMi  iiiillitniiii 

Leptoxis  pliciitii 

Lcptoxis  jinicntsii 

Leptoxis  liiiiiidlii 

Lcptoxis  lir^dtd 

Etimiti  cilabtnucusis 

Elirnia  caclatnra 

Eliiuia  hijdci 

F.Iiiiiiti  olimld 

juffi  silicuhi 

Pleurocera  cdiitiliriiltiliiin  liliiiu 

Plcurorrni  jtiyisiiiiiliiin 

rlrui'iHCi'd  It  iilki'i'i 

Mrliiiioidrs  liiliiiridiilil 


00000 
01000 
00100 
00000 
00000 
00000 
21000 
21002 

?  1  n  0  -• 


00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
noooo 
tOOO 


02010102 
02111012 
02111101 
02111012 
02010102 
02111002 
01101100 
11101100 
11101100 
01101010 

1  r\  1  n  1  1  n  f) 


LU02000UU10111110 


0212101 
0212111 
1002101 
0212102 
0212101 
1212101 
1001111 
1001111 
1002111 
1001101 
3020011 
1011122 
'  "  : 0122 
.1122 

joionj: 


Appendix  4.   (;i'iiiis  l.ilhiisiii    Classification  scliciiirs  used  in  comparison  to  claciistic  li\potlics(>s.  Taxa  marked  until  a  (*)  were  not 
treated  li\  tlie  i>ri<j;mal  .iiitlior  lint  are  included  in  the  primps  liased  mi  tlnir  works  isee  te\(  lor  explanation'. 


Tnr<jeon  et  al. 


Bnrel 


loltl 


I9S() 


Coodneh,   1940 


Tnoii.  I  ST.) 


I)a\is,  1U74 


rein  IS  Litluisid 
L   driiii^ciyi 
L.  riiiid 
L.  (Iiittonidiui 
L   iicniciilatd 
L.  hidirichti 
L.  jnijana 
L.  lima 
L.  oboidtd 
L.  sdlchnisd 
L.  icrmcosd 


(icnns  Lididsid 
.Snlit;eniis  Lilluisid 

L.  flcniciddtd  •j,ciiiciddtd 

L  i^ciiicidiita  fidiiiiiiosd 

L   iicniculiild  piii'j^iiis 

L.  ohoidtd 

L.  snicbvosd  sdlcbrosd 

L   s(drbnisd  florcntidiid 

L   \idrhni\d  sidnjohosd 
Snligenns  Aii<j,i1n-iud 

L   dnni'^crd 

L    rnrtd 

L    diillniudiid 

L    huhnrlili 

L   jdi/dUd 

L    liiiid 

L    nrniciisd 


',r\\\is  l.ilhdsid 

( ironp  1 

/,    iiriiii'J^ri'd 
I.    dllllnilidlld 
I.    Idljaild 
I.    liiiid 

( ironp  2 
/,   'J^i  iiiciildid  <^cniciddtd 
I,    •J,riiiculdld  fidi0nosd 
L.  iifiiicnldld  pin<itiis 
L.  sdlcbrosd  s(dcbrosd 
L.  Sdlcbrosd  jiorcntidiui 
L.  Sdlcbrosd  sul)<slobosd 

Cronp  •■! 
/,   nirld 
L   liiibriiliU* 
I.    irirutosd 

(iroiip   I 
/,    obonilii 


Genus  Ancidosd 

L    '^ciiiciildld  piiiiliiis 
( iemis  Aicj^ilrciiid 
dnni\2.crd 
rnrtd  ■ 

..  llllllolliillld 

..  '^ciiiciildld  '^cuiculdtd 

,   hiilirirlili'- 

.    \d[\dna 

.    liiiid 

,,  sdlcbrosd  sulchrosii 

,.  rcrnicosd 

nns  Lithdsiti 

.   <^cniciddld  jidii^iiiosd 

..  oboidid 

,.  sdli'brosd  fliirciilidiid 

,    sdli'brosd  sidr'loliosd 


(a'nns  lo 

fliiridlis 

iinniiH'rd 

curld 

duttonidud 

'jfuiculatd  ^cniculdtd 

<S.ciiicul(ild  fulifiinosd 

•iciiiciddtd  piii<^iiis 

liiibriihli 

jdt/diid 

liiiiii 

obordia 

sdlcbrosd 

Sdlcbrosd  florcntidiid 

Sdlcbrosd  siib^lobosa 

icrnicosa 


THE  XAITILUS  116(2):50-55.  2002 


Pasrr  50 


CoraIIi()j)Juhi  kaofifonoii,  a  new  species  (Gastropoda: 
Coralliophilidae)  from  the  Canaiy  Islands  Ihing  on  AiitipatJies 
wollastoni  (Cnidaria:  Anthozoa:  Antipatharia) 


Ricardo  N'ega 

Aliapanoii,  65 
29(1  IS  Malaga 
Sl'\l\ 


Rodolfo  Vega 

A\da.  dt'  los  Meiiceses.  38 
3820.5  La  Laguiia  (Tenerife) 
SPAIN 


Angel  A.  Luqiie' 

Lahoratorio  de  Bioiogfa  Marina. 

DfpartanK-iiti)  dc  Bioloiii'a 
Facultad  de  ( Jcncias.  Universidad 

Autoiioiiia 
28049  Madrid 
SPAIN 
angel. lii(jue(S  I  lain. cs 


AHSJKU.T 

.A  iK'w  spt'cius  ol  Cdralliophild  Iroiii  tlie  (,'anan'  Islands  iiviiisi 
on  the  antipatiiariaii  Antijidthcs  udlln.stoni  is  desciilied.  The 
new  speeics  diliers  Iroin  its  closest  Eastern  ,\tlantic  relatiNc 
Coralliopliila  lirciis  iBlaiiuille.  1832)  h\  its  smaller  si/e  and 
spiral  cords  ol  similar  widtli.  and  Irom  the  Western  Atlantic 
Covdlliopliila  ciirihdcii  \l>liiitt.  IV).58.  li\  liavint;  a  thinner  shell 
and  narrower  a[)ertun'  and  In  the  scnlptiirc  and  smaller  size 
ol  the  protoconch.  .Additional  remarks  on  the  t;L\onom\  and 
feetlin<r  of  some  Easleni  .Atlantic  and  Mediterranean  Corallio- 
jihilidae  are  incliiileil. 


IXTKODrcTlOX 

Tlic  laiiiiK  ( '(irallii)pliiliilac  ((iiiipriscs  tropical  tci  tciii- 
pcrate,  siiallow-  to  deep-water  niariiii'  species  tiiat  li\t' 
and  feed  on  cnidarians.  The  coiicliolo^ical  characters 
may  \ai-\-  <freatly  wilhin  the  rainiK.  and  there  ari"  dilTer- 
ent  grades  of  interspecific  and  iiitraspeeifie  \ariatioii 
Shell  \arial)ilit\-,  togetlier  willi  llii'  raril\  ol  most  of  the 
species  due  to  their  restricted  haliitat  (specialK  those 
li\ing  in  deep  water),  the  absence  of  radiila,  tlie  absence 
ol  protoconch.  nsnaliv  eroded  in  adults  and  c\eii  in 
yoiiiig  specimens,  and  the  poor  kiio\\leilij;e  ol  anafoniv 
and  hiol()g\'  have  led  to  a  conliised  taxononix  of  the  fam- 
ily tliat  inaki's  it  tlie  generic  allocation  of  specii's  diffi- 
cnlt.  The  (airalliophilidae  lias  been  IradilionalK  consid- 
ered a  .separali  family  closeK  ri-laled  lo  Muricidac.  but 
Ponder  and  Waivii  i  19SS),  witlioiil  linlhei'  discussion. 
proposed  Corallioplnlinac  as  a  sublamiK  of  Mmieidae. 
This  taxonomic  si  ihis  has  been  adopted  b\  some  siib- 
se(|iicnl  authors  i  Poiidei  l!J9S:  liiedel.  2()()()).  ilowe\er. 
Kantor  ( 1995,  199(Vi  Idniul  enough  differences  in  foregnl 
anatomy  to  jiistif\  tin  ;:  separation  into  two  different 
families.  Since  the  taxonomic  status  of  the  Coralliophil- 
idae is  cnrrentK  nuclei-  diM  nssiou,  uc  prefer  to  use  the 


traditioualK  acceptetl  ranking  ol  tlii'  taxnn  at  the  l.iniiK 
le\'el. 

No  critical  re\isions  ot  the  taniilx'  haw  been  published 
except  for  tliat  of  Vlassin  (1982)  for  the  genera  -A/(/g(7(/.s 
and  Li'ptocoiichus.  but  two  catalogues  (D'.Attilio,  197S; 
Kosuge  and  Suzuki,   1985)  consider  about  200  Recent 


species.  At  least 


<iei 


lera  ha\e  been  d 


lescnhed 


but  the 


Author  for  torrespoTKleiici 


actual  number  is  probabK'  smaller  (DWtfiliu  and  Beitseh. 
1979:  Kosuge  and  Su/nki.  1985';  a  largi'  number  ol  s[ie- 
cies  (about  80)  are  inclucled  in  tlie  genus  CordU'wplnld 
II.  and  .\,  ,\dams.  18.53.  but  this  genus  still  needs  to  be 
better  dehiietl.  .Most  coralliophilids  Iced  on  scleractinian 
corals,  but  there  are  also  spi'cies  that  ti'i'il  on  (ioigonacea. 
C.'orallimoipharia.  Zoaiithidea  (Miller,  1981;  Robertson, 
1970.  1981;  Wicksti'ii  and  Wright,  1993).  Actiniaria  (Spa- 
d;i,  1979;  ()li\erio,  I989;ii.  and  .-Ucvonacea  (Lorenz. 
I99(il  OiiK  h\()  species  ol  Rhizochiius  iRliizocliiliis  an- 
tijuilliinii  Steenstnip.  1850.  and  Rhhi>chihis  sp.'  and  ('or- 
ullii'hid  iiiiiiiii'j^ii  (II.  and  .\.  .\il;ims.  1S6.3)  are  known  to 
Iced  nil  \iitip;illiaria  in  the  Inilo-PaciHc  ( Iva\'.  1979;  Poor- 
iiuin.  1981:  Kosuge  ;ind  Su/uki.  1985;  I^Wttilio  and  Ko- 
suge. 19881  The  t\pe  ol  U'l'diiig  and  the  degree  ol  as- 
soci;ition  willi  :intlio/o;ins  \:u'\  Irom  boring,  eiidobiotic, 
almost  p;ii':isilic.  liigliK  hosl-selecti\e  spi'cies  (.\/r/g(7(/.v. 
LcptdCiiiH-liiis.  Rcl'HjituuTdid)  (Nhissin.  1982.  1983.  1987. 
1988.  1990).  to  less  selective  species  with  a  relati\'el\-  high 
mobiiih  i(',i)v////<7)/i/7r/>  iMiller.   1981). 

Kelerenees  lo  I'liistern  .Atlantic  coralliophilids  are  scat- 
teied  in  ])ublic;itioiis  Irom  dillerenl  ;ire;is:  Meiliterni- 
iiiMii  ;md  I'.iirope.in  ;\ll.iiilie  iS;ibelli  :in(l  Sp:iila.  U)8(): 
()li\eiio.  I989;i.  li;  Poppe  and  (ioto.  1991',  cnnliiieiihil 
WesI  Alric;i  iKmidseii,  1956;  Bernard.  1981;  laiaxeni, 
1975;  (;ol:is.  Piiito-.AIonso  and  Brandao.  19S5:  Kosugi- 
.111(1  Fenumdes.  198S;  Smriglio  :ind  Maiiotliui.  20001. 
deep  w;ilers  ol  the  iioiihtMslerii  All.inlie  i  Honchet  ;iiid 
Waren.  1985).  (.ape  \erde  Iskmds  (Cosel,  1982),  Sao 
Tome  Island  (Kosuge  :ind  Kernandes,  1989:  Rohin  and 
PeriKiiides.  1990),  Saiiil  llcleii.i  Isi.ind  (Smith,  1890), 
;iiid  iVislaii  (!:i('niili;i  IsLuul  Walsoii.   1886).  Four  spi'- 


R.  Wiia  ct  al.,  20(12 


Paei'  5! 


cies  were  recorded  from  the  Canan'  Islands  In-  Nord- 
sieck  and  Garci'a-Talaxera  (1979):  Coralliojiliila  luci/fii- 
cloijfii  (Caleara,  1.S451.  C.  hiviis  (Blainxillc.  \H32}.'  Bti- 
bclomiiivx  cahnifcnis  (G.  B.  Sowerbx;  1834)  las  Coral- 
liophihi  bahclis  (Requien,  1848)),  and  ConiUiophila 
riclwrdi  (P.  Fiselier,  1882)  (as  C.  lacfuca  (Dall,  1889)). 
In  addition  to  C.  richardi.  hvo  additional  deep-sea  spe- 
cies are  known  from  tlie  Canan-  Islands:  C.  sqiuimosa 
(Bivona,  1838)  and  C  Ixisilcus  (Dantzenbersj  and  Fi- 
scher, 1896)  (Boucliet  and  W'aren,  1985).  FinalK",  Cor- 
(lUiophila  foutanaii^ioi/i  Smriglio  and  Mariottini.  2()()(). 
has  been  recenth'  described  from  Tenerife. 

During  SCUBA-cli\ing  samphng  around  the  island  of 
Tenerife  (Canan-  Islands),  a  coralliopliilid  living  on  the 
black  coral  Aiitipatlws  wolla.stoni  (Cra\ )  has  been  found. 
This  third  species  ol  the  famil\-  knou-n  to  li\e  on  anti- 
patharians  is  described  here  as  a  new-  species,  since  it 
shows  clear  differences  from  the  pre\iousl\-  known  cor- 
alliophilids  iroTU  the  .\tlantic  Ocean. 

Institutional  abbresiations  used  in  this  work  are:  BAU, 
Dipartimento  di  Biologia  .'\nimale  e  deU'LJomo,  "La 
Sapienza"  Uni\-ersit\-,  Rome,  ltal\-;  MNCN,  Mnseo  Na- 
cional  de  Ciencias  Naturales,  Madrid,  Spain:  MCNT, 
Museo  de  Ciencias  Naturales  de  Tenerife,  Tenerife, 
Spain;  DBU.\,  Departamento  de  Biologia,  Unixersidad 
Autonoma,  Madrid.  Spain. 

SYSTEM.\TICS 

SuperlamiK  Muricoidea  Rafinesi|ue.  191.5 
FamiK-  Coralliophilidae  Chenu,  1859 
Genus  C(>nilliiij)liilii  II.  and  A.  .\dams.  1853 

Type  species:  Fiisiis  niiitoUlcns  Lamarck,  iSKi,  In 
subsequent  designation  (Iredale,  1912).  Recent.  Indo- 
Pacific. 

Coralliiipliilii  kiuililiumn  new  species 
(Figiu-es  1-7) 

Description:  Shell  i  Figures  1-4'  up  to  24. H  nun  of 
length.  15.5  mm  width,  solid  but  ratlier  thin,  ovoid-iu- 
siifirm:  spiri'  high,  conical,  with  up  to  7  con\e\  whorls, 
shoulder  slightK  angulate:  suture  well  defined  with  un- 
dulate narrow  groove.  Slu'Il  color  uuilorniK  vellowish- 
or  pale-brown,  milk-Ti -white  when  cleaned.  Protocouch 
(Figures  5-7)  of  about  3y2-4  w-horls,  1  uuu  length.  940 
fj-Hi  width.  :ukI  diametei-  ol  first  whoil  280  |jLni.  Pioto- 
conch  indicative  ol  [ilanktotrophic  lai-val  development. 
Protocouch  nucleus  (Figui-e  7i  smooth  except  lor  small, 
scattered  pustules,  subseciuent  whorls  with  tw-o  strong 
spiral  keels,  crossed  b\-  axial  ribs  tormiug  nodules  at  in- 
tersections and  scattered,  small  pustules  (Figure  fS). 
Thick  xarLx  inilicates  limit  between  protocouch  :m(l  te- 
leoconch.  Protocouch  color  pink  to  reddish-  or  pale- 
brown.  Protocouch  lacking  or  usuall\-  eroded  iu  adult 
specimens,  but  when  present  is  frequeutlv  somewhat  lilt- 
ed in  relation  to  shell  axis.  Teleoconch  with  5-7  convex 
whorls,  somewhat  angulate  at  shoulder  on  eai'lier  w  liorls. 
Last  whorl  larsie,  convex,  more  or  less  angulate  at  shoul- 


der, then  uarrowiug  toward  base,  representing  almost  3/ 
4  ol  total  shell  length.  .Aperture  large,  oval,  lirate  and 
white  within:  but  one  specimen  with  inner  side  of  ap- 
erture pinkish.  Outer  li]i  thin,  with  fiuelv-  serrated  outer 
niargin  (margin  rippled  internallv):  inner  lip  slightlv  an- 
gulate in  middle.  Siphonal  canal  broad,  motlerateK- elon- 
gate, open,  slightlv  recuned.  Umbilical  aperture  absent. 
,A\ial  sculpture  of  9-11  ion  specimens  lougei'  than  10 
mm)  or  8-9  (on  smaller  specimens)  broad,  elevated  axial 
ribs  on  last  wliorl,  and  usuallv-  10  axial  ribs  on  penulti- 
mate whorl.  .Axial  ribs  less  marketl  towai'd  end  of  bodv 
whorl  in  longest  specimens,  but  clearlv-  mai-ked  in  small- 
er (younger)  specimens.  Spiral  sculpture  on  last  whorl 
of  up  to  18-19  cords  of  rounded  cross-section,  all  of 
similar  width,  and  bearing  prominent  denselv  packed, 
fine  and  long,  scale-like  lamellae.  Spiral  cords  8-9  on 
the  penultimate  whorl.  Shell  usuallv  covered  bv  encrust- 
ing organisms  [Miiiuicind.  bn-ozoans.  seqiulid  polv- 
chaetes,  two  different  small  species  of  bivalves,  sponges 
and  calcareous  algae  were  obsened  on  a  single  speci- 
men). Living  animal  white  oi- whitish,  inc-luding  tentacles 
and  siphon,  with  deuselv-packed,  vellovv  speckles,  which 
form  a  ring  on  distal  part  of  the  siphon.  Operculum 
brown,  mahogauv-,  or  reddish-brown. 

Type   Material:      llolotvpe  (Figures    1-2  .   .\L\CN" 

15.05/32583.  20.0  mm  length  X  13.5  mm  width:  five 
paratxpes,  .MNCN  15.05/32583:  txvo  paratxpes,  .VICNT 
TFMCBMMO/000277  and  TF.\1CBMM6/()00278:  txvo 
paratxpes,  DBUA.  15.05/18.  All  from  tvpe  localitx-,  at- 
tached to  black  coral  Aiilijiallics  wcillasloiii  (Clrav).  Xov.- 
Dec.  1994, 

Type  Localitx:  Puula  de  Tt-no,  Tenerife.  C^anan  Is- 
lands, 28°20'  N,  17°55'  W.  depth  22-24  m. 

Other  Material  Exaniined:  Twelve  specimens  Irom 
the  txpe  loi'alitv.  Ricardo  and  Hodolfo  \ega  Collection, 
txpe  localitx.  R'.  and  R.  Wga  coll.  Nov.-bec.  1994:  18 
specimens,  B.VU,  Punta  de  Teno:  12  specimens,  BAU, 
Diente  de  .\jo:  21  specimeus.  BAU.  N'eril  de  Masca,  all 
Tenerife,  Canan  Islands.  \l.  Oliverio  coll.:  one  speci- 
men, 20.9  mm  length  x  13.fi  nun  width,  R.  and  R.  X'ega 
Collection,  La  Bombilla.  Palma  Island.  Canan  Islands, 
collected  aliv-e  on  Aniipatlic.s  udlla^lciii.  40  ni  depth.  1). 
R.  Gallowav coll..  24  Aug.  1996. 

Distribution:  known  onlv  Irom  the  tvpe  localitv,  Pun- 
ta de  Teuo.  two  olhei-  uearliv  points  ou  the  Tenerife 
Island  (Diente  de  .\jo  and  N'eril  de  Masca',  and  from  La 
Palma  Island  (La  Bombilla.  David  Roig  (ialloway.  leg.). 
The  planktotro|)hic  tvpe  ol  development  indicated  bv- 
tlie  protocouch  of  CondliopJiila  kaojUonun  suggests  a 
wider  distribution  than  that  currentiv  known,  but  the 
sjiecies  has  not  vet  been  lound  in  other  Maearoncsian 
Islands,  the  West  .\fricau  coast  or  the  Mediterranean. 

Habitat:  .\I1  collected  and  obsened  specimens  vvero" 
found  attached  near  the  base  of  colonies  of  the  black 
coral  Aulipiilhi\  uolldsloiii  (Crav)  or  near  the  major 
branch  nodi's  in  the  case  oi  ven  large  colonies,  between 


Patle  52 


THE  XAUTH.US.  \nl.  11(1  No.  2 


Fifjures  1-7.  i'oidlluiiihdn  kdiifittiniiit  new  sprcics.  I,  2.  llolotxpe,  MNCN,  15.05/32583.  24. (i  X  15.5  nun.  I'linta  (le  Tciio. 
TfiKTife,  Caiian  Islaiul.s.  3,  4.  l\iia(\pc  1.  Mi\t;N,  I5.()5/325S3.  from  the  hpc  Icicalih.  14.9  x  i).fi  mm.  5,  (i,  7.  I'lotouoncli  ol 
paratope  I.  5.  (Iciieral  \i('\\.  Scale  liar:  200  ijliii.  6.  Dt'tail  of  the  sciilptmf  of  the  last  wliorf  Scale  liar;  10(1  jinr  7.  Drtail  of  tlit" 
milieus  ami  tlic  firsi  wliorl.  St-alc  liar:   10(1  \y\n. 


18-30  Ml  iFiinta  dc  Tcmio  ami  Dicnlc  ilr  .\|())  ami  42-4S 
m  (\'eril  de  Masca).  An  averagt'  iiiinilicr  ol  5-6  speci- 
mens per  antipathariaii  colonv  were  ohsencd  in  Pnnta 
de  Teno,  with  a  nuLxiniuin  of  10-12  specimens,  and  np 
t(j  20  on  a  single  antipatliarian  colonx  in  \eril  de  Ma.sca. 
Of  27  .specimens  checked  for  their  posilinn  on  three  col- 
onies, 20  were  with  the  si])hon  directed  upwards  and  7 
do-Anwards  (()|i\erio,  pers.  eonim.).  No  ohservatiuns  of 
(ceding  on  I'ik    I'laek  coral  have  been  made. 

Etymology:  "flic  specific  name  comliines  to  the  nick- 
names ol  the  soii'^  ,i!  tin  first  two  authors:  Kao  (Hicardo) 
and  Ofito  i  Rodollo 

Remarks:  Comllini'hila  kaofilonini  is  tenlalixcK  as- 
signed to  the  genus  C'^uallinplula  liecan,se  it  resemliles 
otiier  species  (C.  cunhara.  (,'  brcvAs)  that  are  presentK 
included  in  that  genus,  jiemiiue  a  generic  revision  of  the 
laniiK. 

Coralliopltila  kaojilnniin  differs  from  its  closest  rela- 


ti\('  ('ordllicjiliilii  lii'ciis  iBlaiusille,  lS.-)2i.  which  is  also 
pi'escnt  in  the  (  ^auaiA  Islands  and  has  lu't'U  collected  at 
the  liase  of  the  \-ellow  (lj>j)ln>i!,or^ia  riiiiiiidlis^  and  red 
(/,.  nilnnimd^  gorgouiaus,  oil  Puerto  de  la  Cnv/..  at  20- 
.'52  m  depth.  ;uid  oil  the  northeiii  side  ol  the  Teno  Pen- 
insula, in  identical  en\  ii(  inmental  conditions,  Coriillic- 
jiliild  lirciis  is  ,1  longer  species  mp  to  .'lO-tO  mm'.  \cr\ 
variable,  espeeialK  in  spire  length  and  shape  ol  the  aji- 
erturc  and  the  siphoual  canal:  the  spiral  sculpture  con- 
sists ol  15  25  (illen  uiarkedK  imbricate,  alternating 
bioad  and  thin  cords:  lrec|ueulK  one  or  two  more  swob 
len  coi'(b  on  the  upper  p;irl  ol  the  spue  acceutu:ile  the 
carene  ol  the  w  liorl.  In  the  .Mediterranean,  ('oi'dlliopliilii 
hici  /s  lives  also  on  iioi'gonians  {Pcirtniiiirircii  cliiimiic- 
Iciiii.  I'.iniiicllii  •■ilricliL  l,(ijilii)^<>r<iia  saniiciilosti)  (Olive- 
rio,  l«S9b:  Po|)pc  and  Coto,  1991). 

('ortillidpltila  riirihricd  .•\bl)()tt,  195S.  from  tin'  West- 
ern .'Mlantic  is  a  similar  species,  but  it  is  somewii.it  solid 
and  has  a  wider  ajierlnre  usuallv   purple  on  the  inside. 


R.  \V>'a  ft  al..  2002 


^ai'e  53 


It  lives  on  a  varieh'  of  antliozoaiis  intlmliu'j;  sclcractiiii- 
ans.  gorgonaceaiis.  z.oantliiileans  and  (x)ralliiii()ipliaria 
(Wells  and  Lalli,  1977:  Miller,  19S1:  De  Jong  and  Con- 
mans,  1988).  The  enibnonie  shell  ol  C.onilliopliihi  r/ir- 
ihaca  was  illustrated  hv  Bandel  (1975),  and  the  piotu- 
eoneh  and  lai"\'al  shell  h\  De  |iing  and  (,'ooinans  ( 1988) 
and  Leal  (1991).  The  prottieuneh  of  f.'.  rnrihrica  has  its 
nucleus  dcnscK  eowred  1>\  small  piistnles.  is  somewhat 
longer  (4.5  wiiorls,  1.2  mm  ieiigtii),  and  has  mon-  pi'om- 
inent  nodules  than  tliat  of  (',  kdnjilnnnit  PrestMitK  mo- 
lecular insestigation  is  in  progress  1)\  Marco  C)li\'erio  to 
assay  the  relationships  of  the  ni-w  species  with  Condlio- 
phila  hrcvis  and  Coralliophila  airihucd. 

The  new  species  also  differs  from  C.nniUiojihilii  mctj- 
endoi-ffii  (Calcara,  1845),  another  littoral  species  of  the 
Canai^v  Islands  wideK  distributed  along  the  .Meiliterra- 
nean  and  Eastern  .\tlantic.  wliich  has  a  longer  (nji  to  40 
nun)  and  more  soliti  shell,  with  5-fi  teleoconch  wlioiis 
with  about  13-15  spiral  cords  antl  8-10  strong  axial  ribs, 
canal  rather  short,  and  umbilical  aperture.  In  the  ('anan 
Islands,  C  incijvmloi-ffti  prt-xs  on  Aiuinon'ui  sidtdhi 
(Perez-Sanchez  and  Moreno-Batet.  1991),  and  in  the 
Mediterranean  the  species  is  known  to  \nr\  on  \hi- 
monia  suIcatiL  Chidocora  ciicspilosa.  anil  BiddHiiplnjllui 
airopaca  (Sabelli  and  Spada.  1980;  C)li\erio,  19S9b), 

The  Atlantic-Mediterranean  ('(irdlliopliilii  sipininosd 
(Bivona,  1838)  has  a  longer  (up  to  40-50  mm)  and  more 
soliil  shell,  with  8-10  iLxial  ribs  and  12-20  major  spiral 
cords,  alternating  with  the  same  number  of  lamellose 
cordlets,  the  siplional  canal  is  of  medium  lengtli  and  tlie 
umbilical  aperture  evident  (Bouchet  .md  Waren,  1985; 
Oli\erio,  1989b).  Corallioplnld  s(iiidiiiiisd  pre\s  on  flor- 
gonacea  or  deep-sea  scleractinians  (Oliserio.  1989b). 
CoraUiophild  luontcrosatoi  (Locard,  1897).  from  Spain, 
and  CordUiophila  pwfundicohi  Haas,  1949,  from  Ber- 
nnula,  should  be  considered  junior  s\non\ins  ol  (',  stjiui- 
niosa,  according,  respectixeh,  to  Bouchet  and  Waren 
(1985),  and  Kosuge  and  Suzuki  (1985). 

Cordlliiiphild  hdsilciis  (Dautzenberg  and  II.  Fischer, 
1896)  reaches  up  to  27  nun,  has  :i  thicker  shell  and 
broader  siplional  canal.  It  is  found  on  the  upper  p;iit  nl 
the  insular  slopes  around  Azores  and  the  (^anan  Islantls 
(Bouchet  and  Waren,  1985);  its  prev/host  is  unknown. 

C.nndliophild  riclidrdi  (P.  Fischer,  1882)  is  a  deep-sea 
(batlnal)  species  tliat  li\es  (and  probabK  pre\s)  on  the 
scleractinians  Madrcponi  ocidatd  and  l.iiplulid  prrtiisd 
in  other  Eiastern  Atlantic  localities  (Bouchet  and  Waren. 
1985;  ()li\erio.  1989b.  and  obsenatious  of  the  tliirtl  au- 
thor). The  shell  of  this  species  is  \en  dillerent  from 
t\'pical  C'ondlidphild.  with  8-10  axial  roli;ite  \arices  in- 
sti'ad  ol  solid  ribs  and  with  relati\el\  lew  spnal  cords 
without  tlu'  characteristic  scaK'  sculpture.  (\indlio}>Uild 
rii-lidrdi  is  included  b\'  Kosuge  and  Suzuki  (  1985)  in  the 
genus  Eiiiczdiiiid  Iredale,  1929,  along  with  the  similar 
Ciinillidjiliild  jdrli  Knudsen.  195(S.  lonnd  hniii  Sierra  Le- 
one to  Angola. 

Comlliophila  foiitdudngioiji  Snuiglio  and  .Mariottini. 
2000,  describeil  from  Teno,  Tenerife  Island,  is  snuiller 
(up  to  7.5  nun)  and  has  a  solid  biconical  shiil.  It  li\es 


on  the  scleractiniau  Madracis  rispcndd  Milue-Ldw;n(ls 
and  llaime.  1850  (Perez-Sanchez  and  .Moreuo-Balet. 
1991;  Smriglio  and  Mariottini.  2000;  authors  obsena- 
tioirs  in  Teneiile).  which  pi'ob;ibl\  is  a  MacaroTiesian  eu- 
tli'mism  (Zibiowins,  1980^.  .According  to  Smriglio  ;uid 
.Mariottini  (2000).  ('.oidlliopliila  l(indldiidii<i,i(>i/i  issimihu' 
to  CaraUioplidd  dlliodii'^iddla  ( !•',.  .\.  Smith,  1890)  li'om 
St.  Helena  Island.  Comlliopliild  ididniuiiddliis  Kosuge 
anti  Fernandes.  1989.  from  S;io  i'om(\  ('oriilliopliild  pd- 

rci  Petneh.   1987.  h l'l(irid;i,  ;uid  I'ordlliopliild  iiirld 

Sowerln;  1894.  from  .Mauritius  Island;  all  of  them  clearK 
diffi'r  from  f  kdofilnniin  b\  the  smaller  size  and  dif- 
ferent shell  sluipc. 

The  remaining;  corallio|)hilid  species  found  in  the  (."a- 
nan  Islands,  Bdhcloiiiiii'cx  ((iiiniffnis  [C.  B.  Sowerbx, 
1834)  is  included  in  a  different  genus  on  the  basis  of 
conchological  and  anatomical  li'atnres  (Kosuge  and  Su- 
zuki. i985;  Hichtei'  and  Luijue,  in  press).  This  sprcies 
pre\s  on  the  scleractiniau  Plii/lldiiiiid  tnoiifliczi  (authors' 
obsenatious).  whereas  in  the  Mediterranean  it  pre\s  on 
('Iddiirnni  iiicspiliKii.  Asli'ciidcs  itdijiiildrts  and  I'oli/ci/- 
dtlius  iniiillcrdc  (()li\c-rio.  1989b.  and  authors'  obsena- 
ti(jns);  ol  these  three  kilter  species,  only  PcdljOjiltluis 
iiiiu'llcrdc  might  possihK  be  lonnd  in  the  ( 'anar\  Islands 
(Zibrowius.  I98()i. 

One  other,  [)robabl\  nntlescribed  littoral  species  of 
coralliophiliti  is  found  in  the  (.'anaiA'  Islands  li\ingon  the 
scleractini;in  Driidroplii/llid  rdiiicd  (from  Lanzarote.  40- 
50  in,  C;usta\()  Perez  Dionis  and  Marco  ()li\erio,  pers. 
comm.).  It  is  a  large  (u])  to  40  mm)  and  solid  shell.  \ri\ 
similar  to  that  iliusti;ited  as  ComUiophdd  (i.  jinii  Irom 
C;abon  b\  Bernard  (1984:  pi.  29,  fig.  120). 

Tlie  remaining  .Mediterranean  species  of  Condliophdd 
are  all  \"en'  dilTerent.  Q'.ondliopUdd  pinicniiHaiid  i.Mon- 
ti-rosato.  lSfi9)  is  solid,  with  10  :ixi:il  ribs  anil  22-24  spi- 
ral cords  with  sm;ill  sc;iles.  This  species  lives  at  the  base 
of  Fdrdiiuiiiccd  cluniidlrdii  :ind  associated  to  ('ondliuiu 
nihniin  (Oliveiio.  19891)'  or  Epizoantlins  arcuacciis 
(Templado  et  ak,  1993).  Cortdllnpliila  saphiac  (.Arailas 
and  Benoit.  I87fi)  has  a  solid,  more  or  less  globose  shell, 
similar  to  Ciiiidliiipliild  sijiuniKisn  in  size,  and  li\es  in 
bottoms  with  strong  conilligeuons  eomponeut:  its  diet  is 
unknown  (Olixerio,  1989b). 

.\mong  the  easti'rn  .Atlantic  species,  (.ondliopliilii  dc- 
(hiiiiiis  (Watson.  1880)  from  Niiihtin^ale  Island  (Tristan 
iki  (!unha'  li;is  a  stroni4  shell,  with  a  high,  scalariform 
spire.  relativi'K  small  aperture  and  umbilicus.  C.ordllio- 
pliild  g//())i  (Dautzenberg.  1891 )  from  the  (."ape  \'erde 
Islands  and  Sao  Tomi>  is  a  smaller  species  (up  to  20  nun), 
with  a  thick  elongate  fusiform  shell  usnalK  eroded  and 
iucnisted,  7-9  broad  axi:il  ribs  ;ind  :ibont  27  s]iiral  cords 
on  the  last  whorl  (Kosuge  aTiil  Fernandes,  1989;  Holan 
and  [•'ernaudes,  1990).  Ccndlioplidd  alldiitica  (F.  A. 
Smith.  1890)  from  St.  ilelen:i  IsLiud  is  :i  solid,  small 
species  (17.2  nun),  with  low  axial  ribs  and  s])iral  cords 
with  weak  sculpture,  ('ondliopli'da  rn/llirost())>ui  F.  A. 
Smith.  1890.  also  from  St.  Helena  is  solid,  sub-rhom- 
boidak  and  has  a  reddish  ajK-rture  and  evident  umbili- 
cus. C'lirdlliiipliild  pdlnnlis  i  Iv  .\.  Smith,  IS90i,  from  St. 


Paire  54 


THF.  \Al  TILLS.  Nol.  1 IH.  NO   2 


Helena,  is  small  ( 10.5  inni).  As  far  as  we  know,  the  three 
latter  species  are  onI\-  known  Irom  t\pe  material,  illus- 
trated and  briefl\'  described  h\  Kosuge  and  Siiziild 
(1985). 

Coralliopliila  kraemmeri  Knudsen.  1956.  from  Nige- 
ria, and  CoraHioj)liihi  marrati  Knudsen.  1956.  from  Li- 
beria, ha\e  sohd,  umbilicated  shells;  the  first  one  has  16 
axial  ribs  and  14  spiral  ridges  on  tlie  body  whorl,  whereas 
C.  marrati  has  rounded  ribs  and  onl\  8  prominent  spiral 
ridges  (Knudsen.  1956). 

Coralliophila  occidentale  Kosuge  and  Fernandes. 
1988,  from  .\ngola  is  a  small  (up  to  9.4  mm)  and  rather 
solid  species.  Coralliophila  adansoni  (Kosuge  and  Fer- 
nandes, 1989)  from  Sao  Tome  is  also  solid  and  has  the 
inner  side  of  the  aperture  denticulate.  FinalK;  Corallio- 
phila knudseni  Smrigho  and  Mariottini,  2000,  from  I\or\' 
Coast  and  Coralliophila  schiottei  Smriglio  and  Mariot- 
tini, 2000,  from  Sierra  Leone,  are  smaller  and  liaxe  bi- 
conical  and  more  sohd  shells  than  Coralliophila  kaofi- 
toruin. 

The  remaining  western  Atlantic  species  of  Corallio- 
phila are  also  different.  Coralliophila  aberrant  (C.  B. 
.Adams.  1850)  has  a  thick,  globose  shell  with  strong  spiral 
cords.  Coralliophila  galea  (Ree\e.  1846)  also  has  a  more 
solid  shell  and  wider  aperture.  Coralliophila  salcbrosa 
H.  and  A.  Adams,  1863.  has  an  o\"ate-fnsiform  shell,  with 
angiilate  spire,  and  Coralliophila  scalarifonnis  (La- 
marck, 1822)  has  a  scalarilorm  profile. 

RECOMMENDATION 

Tiie  host  of  Coralliopliila  kaofitonim.  the  anthipatarian 
Antipalhc.s  wolhi^toiii,  forms  in  the  Canarv  Islands  a 
characteristic  communit)-  on  walls  below  50  m  of  depth, 
but  it  can  be  found  at  sh;illower  depths  (18-24  m)  in 
ca\es  or  overhangs  at  certain  sites  (Perez-Sanchez  and 
Moreno-Batet,  1991 ).  A  similar  itssemblage  occurs  in  the 
Caribbean  islands  with  other  antipathaiian  species  (Au- 
tipathcs  spp.,  Humann,  1993).  Antipadies  uollasloni  is 
only  known  with  certaint)  from  Madeira.  Selvageni  and 
Canan  Islands,  but  it  is  probabK'  also  present  in  the 
Cape  \'erde  Lslands  (A.  Brito,  pers.  comm.).  It  is  pro- 
posed for  protection  in  the  Canar\  Islands  and  included 
in  tiie  Prcliniinarv-  Red  List  (Bacaliado  et  al.,  1989)  and 
the  Threatened  Marine  Fauna  List  (Boimet-Femandez- 
Tnijiiio  and  Rodriguez-Fernandez,  1992).  We  strongK 
suggest  to  the  environmental  authorities  of  the  Canan' 
Islands  C()\eniinent  to  include  Coralliophila  kaofitonim 
in  the  li',!  (li  jiriiletttd  species,  since  it  is  up  to  now  onK 
kn(jwn  horn  ili.  (  aiiarv  Islantls  and  is  strictK  associated 
wiih  Antij)iitUi-.  :loII(i.sI<iiu. 

ACKNOW  LF.Dr;MK\TS 

We  are  indebted  l<i  Marco  Olixerio  for  information 
about  the  habitat  of  C.  knofitomm.  and  to  him  and  Al- 
exandra Ricliter  for  in  Iplni  comments  on  the  manu- 
.script:  to  Gu.stavo  Pen/  Dionis  for  data  on  coralliopliii- 
ids  from  the  Canarx   IshnKK;  to  Jose  Bedova  uMNC.N) 


for  SEM  photographs,  and  to  Rogelio  Sanchez  and  Jesiis 
Mufioz  iMNCN)  lor  photographs:  to  Alberto  Brito  and 
Rogeho  Herrera  for  information  about  conserxation. 
This  work  was  supported  h\  Fauna  Iberica  W  project 
(DC1C\T  PB95-0235>. 

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Talavera.  F.  G.  1975.  Molnscos  de  sedimentos  de  la  plataforma 
continental  de  Mauritania.  Boletiu  del  Institute)  Espafiol 
de  Oceaiioijrali'a  192:  1-18. 

Teinplado.  |  .  .\.  Guerra.  J.  Bedma,  D.  Moreno.  J  M.  Remon. 
M.  Maldonado  and  M.  .\.  Ramos.  1993.  Fauna  iiuuina 
circalitoral  del  snr  de  la  Peni'nsula  Iberica.  Resultados  de 
la  Campaua  Oceanognilica  "Fauna  I ".  .Museo  Nacional  de 
Ciencias  Naturales,  Consc^jo  Superior  de  Investigaciones 
Cientfficas,  Madrid.  138  pp..  45  pis. 

Watson.  R.  B.  1886.  Rejiort  on  the  Scaphopoda  and  Ciitstro- 
poda  collected  In  H.  M.  S.  Challenger  during  the  \t^ars 
187:3-76.  Report  of  the  Ciialleiiger  Expedition,  Zoology 
15(42):  756  pp..  50  pis. 

Wells.  F.  E.  and  C.  M  Lalli.  1977.  Reproduction  and  brood 
protection  in  the  Caribbean  gastropods  Coralliophila  ah- 
breviata  and  C.  raribaea.  )ounial  of  Moiluscan  Studies  43: 
79-87. 

Wicksteu,  M.  K  and  K  T  Wright.  1993.  Predation  h\  Latiaxis 
oldroifdi  (Gastrojioda:  Coralliopliilidae)  on  Con/nactis cal- 
ifoniira  (.\ntlio/.oa:  Corallimoqihidae).  The  \'eliger:36:  92. 

Zibrowius.  H.  1980.  I^>s  Sderactiuiaires  de  la  Mediterranee  et 
de  r.Mlantiijne  nord-oriental.  Memoires  de  I'lnstitut 
()ceanos;raplii(iue.  Monaco  II:  1-227. 


THE  NAUTILUS  116(2):56-5.S,  2(){)2 


Patre  56 


Fusi}ms  dovpch'di,  a  new  species  (Gastropoda:  Fasciolariidae) 
from  the  Red  Sea,  and  range  extension  for  two  other  species 


Martin  Avei-)'  Snyder' 

Departiin'ut  ol  Midacologx 
Academy  of  Natural  Sciences 
19'''  and  Beiijaniiii  Franklin  Parkway 
Piiiladeiphia.' I'A  19103  USA 


ABSTRACT 

Fusinu.s  dovpcledi  new  species  from  the  Red  Sea  is  descrihed. 
The  new  species  is  distinguished  hv  shell  characters  from  the 
similar  Red  Sea  species  F.  leptorht/iicliua  (Tapparone-fJanetri, 
IS75)  and  from  F.  coins  lon^caiida  (Lamarck,  ISOl)  from 
southeastern  Africa.  New  range  records  are  provided  for  F. 
hifmns  iSUudm.  1900)  and  F  aralnnis  (MeK-ilf  1898). 


INTKODUCTIOX 


tni 


A  new  species  in  the  li;cihis  I'lishius  is  ilcscrilied  ticiiii 
the  Red  Sea  aiici  tiLxoiioinic  affinities  are  cliscussed.  Most 
Fiisiniis  are  subtidal  burrowers,  although  some  species 
are  tound  in  deep  water.  Tlie  discoNciv  and  description 
ol  this  new  species  is  part  of  a  trend  in  recent  wars  in 
which  a  nnniber  of  new  fasciolariids  have  been  nanietl 
(e.g.,  Gofas,  2000;  Hadom  and  Rogers,  2000;  Snyder, 
2000;  Snyder  and  Snvder,  1999;  and  others).  Specimens 
ol  the  new  species  described  herein  ha\e  been  collected 
by  fishermen  at  a  depth  of  .300  m.  AdditionaJK.  the  re- 
discoverv-  of  Fusiiiii\  hifrciis  iStunuiv.  1900).  another 
Red  Sea  species,  with  a  ran<j;e  e\lension  lo  the  Indian 
Ocean,  is  reported.  ,\  probable  range  extension  is  also 
reported  for  Ftisiniis  anihicus  I  VleKill,  1S9S).  Tin-  ab- 
breviation USNM,  National  Mu.seum  of  Natural  lliston. 
Smitlisonian  institution,  W'asliiii'^lon  DC.  is  used  here. 


SYSTEMATICS 

Family  Fasciolariidae  Gray,  1853 
Snbfaniil)  Fusininae  Wriglev,  1927 
(^i-uns  Fusinns  Rafines(|ue.  IS15 

'Ivpe  .species:     Murex  coins   Linnaeus,    17.5S.   Hecent. 
hidu  \'v.  ~:  Hacific.  by  monot\-py. 

Ftisiiiiis  ildijtclcdi  new  species 
(Figures  1    2' 

Descriplion:      Sh  H  elongate,  .si/e  moderate  lor  genus, 
length  to  107  imn.  Mplional  canal  long,  slightK  nndnlal 

'  Research  Associat* 


ed,  apex  subtending  an  an<j;le  ol  ;ippro.\iniateK-  29\ 
Sculptiu'e  of  iLxi;il  libs  and  spir;il  cords,  ribs  lorming 
prominent  angular  knobs  on  later  whorls,  with  knobs 
hilling  approximateh  at  the  centi'r  ol  whorl.  V>ud\  whorl 
with  11-13  ri])s,  penultimate  and  earl\  wliorls  with  12- 
14  ribs.  Earlv  whorls  ofparatype  (onlv)  lightly  sculptured 
with  ;Lxial  ribs,  becoming  evanescent.  Axial  ribs  crossed 
1)\  unuierous  spiral  cords,  some  lirowii  on  later  whorls; 
strong  brown  cords  alternate  with  weaker  unctjlored 
cords  on  bodv  whorl.  Penultimate  whorl  witli  12  strong 
cords,  becoming  evanescent  toward  the  suture,  luid  un- 
uierous weak,  fine,  axial  lines,  visible  onlv  under  mag- 
iiificiition,  between  axial  libs.  Aperture  tvpicallv  ovate 
;iud  elongate;  parietal  shield  shiny,  waxy  white,  extending 
onto  siphonal  canal.  Spiral  cords  bene;ith  shield  raised 
to  lorin  lirate  sculpture.  Canal  long.  thin,  somewhat  sin- 
uous. Operculum  claw-like,  light  brown.  Piotocoucli. 
periostnicum.  and  nulula  unknown. 

Type  male-rial:  llololxpe  USNM  90.3651.  length  107 
mm,  width  24.4  mm,  spire  height  40  mm,  live-collected. 
P;u'at\pe,  IVled  ( Collection  (d;un;igi'd),  length  72. S  nun, 
width  22. S  unn,  spire  ;iiid  siphon. il  can.il  broken,  dead- 
collected.  Ii'oui  t\pe  loc;ilit\. 

Tvpe  loealilv:  f-led  Se;i  oil  Neviot  (Egvpt),  75  km 
soutli  ol  F.hit  (Israel),  .300  m  depth  on  s;nidv  bottom. 

Etvinologv:  The  species  is  named  ;ilter  l)o\  P<'le(l 
(Tivon,  Israt'l),  a  student,  colleetoi'  and  dealei'  in  the 
shells  ol  the  lU'd  Se;i  ;u('a. 


Distriliulion  and  liabilat: 

c'alilv. 


Know  u  onl\    li'om  t\pe  lo 


Diseussion:  I'nsiiiiis  (hnpclcdi  is  most  closely  rekited 
lo  i  hplnihijuchus  (T;ipp;u'one-( 'auelri.  1875').  which  is 
well  illustrated  ni  Shanib;iti  (19S1:  pf  23,  fig.  7).  The 
knobs  on  F.  Icplorln/iirliiis.  loi'med  b\  the  ;Lxial  ribs,  are 
lewei',  immbeiing  just  9,  ;ind  ;ire  uioi'e  S(|uare  th;m  those 
on  I  (liii  jHlcdi.  The  opi'iculnni  ol  /'  Icplorhtjnchns  is 
distinetivek  reddish  brown.  ,uid  the  shell  is  sm;iller.  tvp- 
ic;dl\  r;iiiging  between  7.5  and  SO  unii.  with  ;i  propor- 
tioiKitek  shortei'  siphon;il  c;mal.  These  dillerences  ;ire 
snllicient  to  iide  out  F.  dovpvlrdi  Ixing  a  deeper-water 


M.  A.  Sinck-r,  2002 


Page  57 


Figures  1-4.  Species  of /•//,s(i///,s,  1,  2.  /■//s;//((.s  doipchdt  neu  species.  I.  Ihilohpe.  I'SNM  U()3(i51.  107  iniii  leii<;tli.  tioiii  saiuK 
hottnni  off  Neviot  (Eg\pt).  75  km  sontli  of  Elat  (Israel),  Ued  Sea.  deptli  300  in.  2.  I>arahpe.  Peled  Collection.  72.8  mm  leiisitli. 
from  hpe  localiU.  3.  Fusinufi  bifrons  (Sturam-.  1900),  Peled  Collection.  99.9  mm  length,  from  Elat,  Red  Sea  (Israel),  depth  .3.50- 
400  m.  4.  Fnsinm  arabirus  (MeKill,  1898).  Snyder  Collection.  96.6  mm  length,  trawled  hy  fishermen  oil  Kl  Tnr,  Gulf  ol  Suez 
(Egvpt),  depth  32^0  in 


Paee  58 


tup:  XAITILUS.  \ol.  llfi.  \< 


lonii  of  F  1c))lorhi/nrlnis.  Comparison  can  also  l)t'  made 
with  /'"  coins  hmgicnuda  (Lamarck,  1801)  troni  the  east 
coast  of  southern  Africa  (Cernoliorsk)',  1972:  pi.  4S,  fig. 
la);  that  species  has  10  axial  ribs  on  the  penultimate 
whorl.  Ho\ve\er,  whereas  the  luunber  of  axial  ribs  in- 
creases toward  the  posterior  end  of  the  spire  of  F.  clo- 
ipeh'di.  the  opposite  is  tnie  in  F.  rolus  hmisicauda.  The 
axial  ribs  are  not  neark  so  pronounced  on  tlie  bodv 
whorl  in  F.  coins  hmiijcaiida  as  in  F.  dorpclcdi;  the  for- 
mer grows  to  appro.ximateh'  200  nun. 

Fiisimis  hifrons  (Sturanx;  1900) 
(Figure  3) 

.Sturan\'  (1900:197)  introduced  a  deep-water  species,  Fn- 
siis  hifrons,  dredged  in  depths  of  490-900  m  in  die  Red 
Sea.  He  compared  this  axialK'  ribbed  species  to  F.  for- 
ceps (Perr\',  1811),  F.  multicarinatns  (Lamarck,  1822),  F. 
torenma  (Desha\'es,  1843)  (a  sviionvm  of  F  coins  (Lin- 
naeus, 1758)),  and  F.  leptorhijncns  (Tapparone-Canefri, 
1875),  and  he  named  a  smooth  varietv'  of  his  new  species 
"form  pancicostata."  Later,  Sturan\'  (1903:  pi.  1)  illus- 
trated F.  bifrons  and  erroneoush^  referred  to  the  taxon 
pancicostata  at  specific  rank.  According  to  R.  Janssen  of 
the  Senckenberg  Museum,  Frankfurt  (reported  prixatelv 
by  R.  Hadorn).  the  t\pe  material  of  this  "form"  differs 
from  that  of  the  nominate  species  onK'  in  sculpture  and 
this  "form"  should  be  considered  a  synonym  of  F.  bij- 
rons. 

Two  IreshK-collected  specimens  of  the  smooth  form 
of  F.  bifrons  were  examined,  one  99.9  mm  long,  26.fi 
mm  width,  with  broken  siphonal  canal  (Peled  Collection, 
from  the  Red  Sea),  the  other  106.5  nun  long,  26.9  nuu 
width  (Sn\der  (Collection,  from  the  Indian  Ocean).  The 
smaller  specimen  was  dredged  on  a  sand\^  bottom  at 
.35()-4()()  m  off  Neviot  (Eg\pt),  75  km  soutli  of  Flat  (Is- 
rael), Red  Sea.  The  larger  specimen  was  collected  by 
Russian  research  expeditions  to  the  Sava  de  Malha  Bank 
in  the  Indian  Ocean  (Bondare\  and  Roeckel,  1992;  Sir- 
(Miko,  1995).  The  larger  specimen  was  illustrated  b\'  Sir- 
enko  (1995:  fig.  7).  This  represents  a  significant  range 
extension  for  this  species  from  the  lied  Se:i  lo  the  indi;ui 
Ocean. 


Fnsinns  anihicns 

(Figure  4i 


McK 


1898) 


.■\  siL'iii'icMiii  r  inge  extension  of  Fnsinns  arabicns  (Mel- 
sill     1S9S-    IS    ;ilsu    reported.   This   species,   previously 


known!  from  the  Culf  of  .Arabia,  is  figured  in  Bosch  et 
al.  (1995:  136,  species  578).  Two  specimens  were  trawled 
by  fishermen  in  the  19S0s  off  El  Tur,  Gulf  of  Suez 
(Eg\pt),  in  depths  of  32—40  m.  Both  specimens  were 
trawled  as  dead  shells  and  hence  this  range  e.xtension  is 
stated  without  absolute  certaintx'.  E\en  though  taken  In 
commercial  trawlers,  with  notoriousK'  xasjue  collectinsi 
data,  it  is  \irtuall\-  certain  that  the  specimens  were  col- 
lected in  the  Red  Sea  rather  than  the  Arabian  Sea.  These 
two  dead-collected  specimens  are  identical  in  ;ilmost  all 
respects  to  Arabian  Sea  specimens  of  F.  arabicns  (Mel- 
vill,  1898). 

LITERATURE  CITED 

Bondarew  I.  and  D.  Roeckel.  f992.  The  shells  of  Sava  de  Mal- 
ha Bank.  La  Conchicrlia  262:  21-38. 

Bosch.  D.  T.  S.  P.  Dance,  R.  G.  .Moolenbeek  and  P  G.  01i\en 
1995.  Seashells  of  eastern  Arabia.  -Motivate  Press.  Dub;u, 
296  pp.,  col.  pis. 

Cernohorsk).  W.  O.  1972.  .Marine  Shells  of  the  Pacific,  II.  Pa- 
cific Publications,  Sydney:  411  pp.,  68  pis. 

Gofas,  S.  2000.  Four  species  of  the  familv  Fasciolariidae  (Gas- 
tropoda)  from  the  north  .\tlantic  seamounts.  loumal  of 
ConchologN'  37:  7-16.  figs.  1  —  3. 

Hadorn.  R.  and  B.  Rogers.  2000.  Re\ision  of  recent  Fusinus 
(Gastropoda:  Fasciolariidae)  from  tropical  western  .Atlan- 
tic, with  description  of  six  new  species.  .Ar^onauta  14(1): 
5-57,  pis.  1-16. 

Sharabati.  D.  1984.  Red  Sea  Shells.  Routlcdsie  6c  Kegan  Paul. 
London:  12S  pp.,  49  pis. 

Sirenko,  B.  I.  1995.  On  the  fauna  oi  shell-bearinij  molluscs  in 
the  Sa\a  de  Malha  Bank.  Indian  Ocean  (part  2i.  La  C^on- 
chiglia  276:  20-24,  figs.  1-15. 

Sn\der.  .\1.  .A.  20()().  Latiiiis  licckt/cc.  a  new  species  ol  F"ascio- 
lariidae  (Gastropoda:  NeoCTastropoda'  Irom  Bra/il.  Tin- 
Nautilus  114:  161-163.  figs';  1-3. 

Snxder,  M.  A.  and  N.  C.  Snyder.  1999.  A  new  species  from 
Columbia.  South  America:  Fiisiiuis  cohunhicnsis  n.sp. 
(Ga.stropoda:  Fasciolariidae  I  La  Conchiglia  291:  21-22, 
figs.  1-1. 

Sturans,  H.  19110.  Diagiioscn  iieuer  Ciastropotlcn  aiis  dem 
Rotlien  ,\leere.  .\n/.eiger  der  Kaiserlichen  .Akadeniie  der 
Wissenschaiten.  Matlieniatisch-NatunNisscnsciiaftliciu' 
Classe  37(17):  197-201. 

Sturaii\,  H.  1903.  Gastropoden  des  Rotlien  Meeres.  Bericlite 
der  (Commission  fiir  Oceanographisclie  Forschungen.  Fx- 
peditionen  S.  M.  Sciiill.  Pola'  in  das  Rotlie  Meer  ncirdli- 
clie  nnd  siidliche  lliilltc  l.S9.5/9r>-I,S97/9S.  Zoologisclie 
Ergebnisst"  23.  Denkschriften  der  K:iiserli<iien  Akadeniic 
der  Wissenschaiten.  .Matheinatiscli-N'atni'wissensiiialdi- 
che  Cla.s,se  74:  209-2S3.  pK.  1-7,  1  text  fig,  (.Mso  issued 
as  a  separate,  pp.  1-75). 


TIIK  NAUTILUS  1  l(S(2);59-fil.  2002 


Viv'v  59 


The  systematic  position  of  Str()]nbi)ia  (Cotonopsis)  J'nidae 
Petuch,  19SS  (Gastropoda:  Columbellidae) 


Helena  Fortunato 

('niter  tor  Tropical  I'lileot'ColoaA  anil 

Arclu'i)I()ti;\ 
SinitliSDiiian  Tropical  Rcscarcli 

Institute 
r,  ().  H(i\  \m 
Balliiia.  PANAMA 
lortiinae(S'ancon. si.edu 


ABSTRACT 

The  s\steniatic  position  of  Stroiiilnim  limliic  I'clneli  I9SS,  is 
discussed  and  revised.  The  species  sliould  lie  allocated  in  the 
colunihellid  genus  Cotonopsis  Olsson.  1^)12  The  shell  char- 
acteristics that  define  the  two  known  Cotoiiojisis  subgenera.  (.'. 
(Cotonopsis)  and  C.  {Tiirrind).  are  iteiui/ed  and  contrasted. 
The  holot\pe  of  S.  liiulac  is  re-illnstrated  and  the  pmtocouch 
Hiiured  tor  tlu'  first  time. 


INTRODUCTION 

111  his  rcNlsion  of  the  Stroniliiiia  o;r()up,  Jtiny;  (19S9)  gaxe 
Cotonopsis  Olsson.  1942.  t^eneric  status  ami  recogiii/ed 
two  subgenera,  Cotonojjsis  scusii  stricio  Olsson.  1942, 
and  Cotonopsis  {Tiiniua)  Jung,  19S9.  Cotonopsis  is  ivp- 
fesented  bv  18  species,  two  of  them  known  only  as  fos- 
sils, and  16  extant.  The  majoritx  of  the  living  species  (12) 
are  found  in  the  eastern  Pacific.  Two  species  are  loinid 
in  the  Caribbean  Sea  (Houbrick.  I9S3:  Fetncli.  I9.S8): 
one  species  in  West  Africa  (Emerson,  1993);  and  a 
fourth  species  in  the  Andaman  Sea  (Kosnge,  Rous.sy  and 
Mnaiiginaii.  1998:  Kronenberg  and  Dekker.  1998,  1999). 
.\s  noted  b\  Kronenberg  and  Dekker  (1998).  this  distri- 
bution miglit  indicate  an  earlier  origin  ol  this  geims.  i.i'. 
prior  to  eaiK  Pliocene  as  proposed  by  Jung  ( 19S9).  On 
the  other  hand,  bodi  the  West  African  anil  the  Asian 
species  ina\"  be  part  of  separate  lineages,  the  laiik  ol 
which  can  onl\  be  addresseil  pcniliiig  aiiatoiiiical  and 
molecular  data. 

The  aims  ol  this  paper  are  to  correct  the  svstematic 
]iositioii  ni  Stroinhina  iColonojisis^  limlur  Petuch.  19SS. 
to  discuss  the  subgeneric  position  ol  tins  species,  and  to 
rc-illn.stratc  the  holotvpe  deposited  in  llie  National  .Mu- 
seum of  Natural  Historv.  Smitlisoiiian  Institution 
(USNM  859942).  with  special  attentimi  to  llir  proto- 
conch.  This  latter  task  was  prompted  b\  bntli  die  poor 
ijualitv  ol  the  original  illustrations  and  the  lack  ol  iiiior- 


iiiatinii  nil  the  1 1 lorpl lologv  ill  the  pi otoci inch  III  tile  orig- 
inal description. 

syst1':m.\tk:s 

FaiiiiK  (  Jiiiiiiilii'llidae  Swaiiisoii.   I S4() 
Cienus  Cotoiiojisi\  ( )lssi)ii.   1942 

Coloiiiipsi^  oKsoii.  Ii)l2:  227  i7o'.  T\pe  sjiecics  (In- original 
desi^iiatiuii  1;  SI  ro  nihil  III  Coloiiojisis  •  iiiiiuicostiirii-cnsis 
OKsoii.    1042  lOlsson.   I!)t2:  7."x  pi,    1(1.  fig.   10'. 

Deseriplioii:  Shell  small  to  large  >  1  l-5(S  iimiL  general 
shell  shape  \ar\ing  Iroiii  lairK  stout  to  slender.  Most 
species  lightlv  sculptured  with  axial  ribs  predominantly 
on  earK  teleneniiili  whorls.  Outer  lip  usnallv  wcaklv 
thickeueil.  both  cohiiucllar  and  parietal  callus  usually 
present,  weakK  to  well  developed.  Parietal  ridge  and 
posterior  canal  present,  prominent  in  several  species. 
Reclined  and  well-developed  anterior  canal.  Proto- 
coucli  smooth  with  I'i  to  :5  whorls,  i  according  to  Jung, 
19SS.) 


Siibgeiiiis  Colonopsi\  sinsn  sirioto 

Description:  Sti.iit  shells,  inllateil  bodv  vvlioH.  axial  ribs 
on  earlv  teleoioiieh.  well  developed  parietal  and  colu- 
mellar  callus,  weaklv  developed  ixirietal  ridge  and  pos- 
terior canal,  and  smooth  protocoiieli  with  I  ':  to  3 
w  liorls. 

Cotinnijisis  (Cotonopsis^  lindiir  (I'etiieh.  I9SS1  new 

eombination 

(Figures  1-6) 

SlioiiibiiKi  tCotonopsisi  lindfir  Petuch.  199S:  161-lfi2.  pi.  .3S. 
figs.    15-16   (Off  St.   James,    Badiados.   trawled   TO   in 

depth,  1 

Descriplion:  Shell  small,  stout,  up  to  22.9  mm  in 
length,  mllated  bodv  whorl,  axial  ribs  on  the  eariv  teleo- 
com-li.  relativelv well  developed  colimiellar  and  parietal 


Page  HO 


TUF,  NAUTILUS,  Yd.  116,  \( 


1^ 


Figures  1—4 

l,atenil  \ic\\. 


:>'^ 


('(>li>U(>ii\is  {('(tidiiojisi.',!  liiiddc    lioldhpc.  I'SWI  S.i!)iM2,  lieiij;lit  22. SS  iiiiii,  wultli  VJ.dO  mm.  1.  .\pcrtuml  \ic-\\.  2. 
ri^lil  siili'    3.    Miapcitiiia!  \ic\\.  4.  I'mtdciiiuii  aiul  caiK    Iclcdrniii-li  wlidils. 


calluses.  I'lolocDucli  siikioIIi,  witli  154  wliorl.s  and  .ilicnil 
TOO  fiiii  iliaiiiclcr. 

Discussion:  'I'lic  hvo  siil)y;cncra  rccocriiizcd  In'  Jmii!; 
!  HISS  I  (lillcr  ill  several  inoipliological  Iratiircs.  In  con- 
trast to  the  ciiaractcrs  altoxc  dcscrilicd  lor  ('dtdiinjisis 
sciisu  slrirto,  species  ol  Cotoiidiisis  {'runiiia)  have 
smooth.  niostl\-  slender  sliells,  with  well-  and  sometimes 
heavily  (le\(>l()ped  pari(Hal  ridge  and  posleiioi  laiiai;  pa- 
rietal and  colnniellar  callnses  missing  or  vcia  weak,  and 
a  smooth  protoconeh  with  2  %  to  3  whorls.  Based  on 
shell  eiiaracteristics.  C.  liudac  is  host  placed  ni  die  snh- 
genus  C(>li)iii>j).sis  scnstt  slrirfo. 

Colotiopfiis  {Titniua)  seems  lo  he  reslncled  to  (lie 
eastern  I'aeific  aTid  has  no  known  fossil  reconl.  ('dlnii- 
(tpsis  sciisu  stricio  has  a  much  wider  disdilmlion,  in- 
eliuling  llie  eastern  Pacific,  the  Carihlx-an  .Sea.  the  West 
■\tri(  an  coast  and  the  .\ndainan  .Sea.  StratigraphiealK. 
the  latter  ranges  from  the  earK  Pliocene  ol'  l'",smeraldas, 
Keuador.  .ind  ( "harco  \vw\.  Peninsula  Burica,  Costa  Hiea, 
through  (he  Rei  ent.  As  mentioned  ahoxc,  the  recent  dis- 
tril)utio;i  ■)!  th.is  snljgeims  seems  to  indicate  an  origin 
older  than  (he  I'iinr cni'. 
This  is  the  sni.illi  St   of  all   lour  species   repoited   li-oni 


outside  (he  eas(ern  Pacific,  ('(iloiinjisis  liiiil/ii'  resemhles 
(,'.  i>liukclc)isis  (Kosnge,  Houss\'  and  .\Inangman.  1998) 
Ironi  the  \ndanian  Sea  in  its  reduced  sculptural  ele- 
ments, hut  the  la((ei"  is  .i  mueli  kirger  .species,  with  an 
incised  suture  and  a  protoconeh  with  a  larger  number 
ol  whorls.  Cotoiiopsis  fin^ciitcfi  llouhrick.  19S.3,  from  the 
Dcinnuican  Hepulilic  and  (.'ii/iiiio/jxi.s  monfilsi  Emerson, 
lOO.i.  Ironi  Si'uegal  are  ,ilso  unich  larger,  with  hea\ier 
sculptured  teleocoucli  whoils.  Besides.  ( '.  inoujilsi  lacks 
axial  rihs,  hearing  insteatl  sjiiral  lines.  Its  protoconeh  has 
(hree  whorls.  The  ])rotoconch  ol  C.  ar<irnlc(i  is  similar 
(o  (hat  ol  ( ',  liiidiic  in  si/.e.  lin(  the  lornier  has  a  weaker 
colnmell.ii  (■.liins. 

.\mon'4  till'  eastern  I'aeilie  species,  (he  diiiiensioiis  ol 
('  lindiic  are  similai'  (o  Coloiiiipsis  iitcndozaiKl  iSli,isk\. 
1970^  from  the  (  aiH' of  h'onseea,  Kl  SaKador.  lint  (he 
i.ittei  h.is  .1  iiineh  less  reciiiAed  siplioiial  ean.il.  more 
liea\ll\  senlplnred  e.ilK  teleocoucli  wliiirls.  \\ell-de\el- 
oped  eiilnniell.ii  callus,  and  ineiinspicuDiis  liiae  on  ils 
ou(er  lip.  SimilaiK  (o  ('  liiidiic.  ('()li»uipsis  isincraldcit- 
sis  (Olsson.  19(iU  Ironi  the  I'.smeraldas  pro\ince,  I'x-- 
iiador.  also  lacks  lieaw  seiilptme  and  li.is  alinost  (he 
same  dimensions:  |iii(  i(s  siphonal  canal  is  lnn'j;ei  ,nid  it 
has  a  iiioic  strongK    dexeloped  eoliiinellai"  callus 


11,  Fortiiiiatd.  2(102 


Fauc  61 


Figures  5-6.  Ci>tono))sis  (Cotonopsis)  lindac.  liolotNpc, 
USXM  S59942.  5.  SEM  of  protoconeh,  sliowins;  a  slijj;litl\  sig- 
moid !ip.  and  the  sculptuie  of  tlu'  carK'  tt'lcocoiit-li  wluirls.  6. 
Same,  apical  \ie\v.  Sade  lines  =  200  (xni. 


ACKXOWI.KDCNIKXTS 


Special  tliaiiks  arc  due  to  |ern-  llarasewvcli  (USWl) 
who  kiii<ll\  photoiiraphed  the  liol<)t\pe  and  helped  in 
main  other  \\a\s.  Tlianks  also  to  two  anoininous  re\ ■Jew- 
els who  kindl\'  read  the  pajicr  and  made  helpful  sug- 
"estioiis. 


LITERATI  RE  CITEU 


Emerson.  \\.  K,  KW'v  .\  new  species  of  coluinhcllid  gastropod 
from  the  old  world  tropics.  The  Nautilus  106:  147-151. 

Houhrick.  f-i.  S.  19S.">.  .\  new  Stromhiiia  species  (Gastropoda: 
I'rosohranchiai  from  The  tropical  Western  .Atlantic.  I'ro- 
ceedinsis  of  the  Biological  Societ\  of  \\'ashin<iton  96:  3-19- 
354. 

Jung,  P.  19S9.  Re\isioii  of  the  Slnmihinti-(.',nmp  (Gastropoda: 
Columbellidae),  fossil  and  li\ing.  Distribution,  biostratig- 
raphv  and  svstematics.  Mcinoires  Suisses  de  Paleontology 

in:'l-29S.' 

IvoMiiie.  ,S,,  P.  il,  Hoiiss\,  and  I'-i^,  Mnangman.  199S.  f^eport 
on  the  fauna  of  Thailand  ( 1  i  with  the  description  of  a  new 
species  ((Columbellidae  and  Ikiccinidae).  Bulletin  of  the 
Institute  of  Malacolog\  ol  Tok\o  3(5):  75-76. 

fCronenbcrg,  G.  C.  and  II.  Dekker.  1998.  A  new  .species  of 
Colonops!.\  Olsson,  1942,  from  an  unexpected  l()caljt\ 
(Gastropoda  Prosohr.iiK  hi.i:  (Columbellidae'.  X'ita  .Marina 
45(3-4):  11-16. 

Kronenberg.  G.  G,  and  II,  Dekker  1999.  ('Dloiiiipsis  viiiintil- 
Ic'^liciiii  Kronenberg  i^  Dekker  199S.  a  junior  s\iion\in  of 
Siroiiihiiiii  phukclcii\i.s  Kosuge.  Kousss  6:  Mnangman. 
I99S,  with  som(>  notes  on  the  generic  position  and  colour 
pattern  i  GiLstropoda:  Pro.sobranchia:  Golumbdhdae).  \'ita 
Manila  46(1-2):  69-72. 

(Olsson.  .v.  .\.  1942.  Tertian  and  Quaternary  fossils  from  the 
Burica  Peninsula  of  Panama  and  (.'osta  Hica.  Bulletins  ol 
American  Paleoutologx  27  if06):  f53-25S  if-I()6!.  pis. 
14-25(1-121. 

Petiich.  E,  ].  19SS.  Neoneuc  hislon  oi  tropical  ,\merican  niol- 
Insks.  The  Coastal  Education  6c  Research  Foundation 
(GERE),  Charlottesville,  1-217  pp. 


THE  NAUTILUS  1 16(2):62-65.  2002 


Page  62 


The  reproducthe  anatomy,  taxonomic  status,  and  range  of 
Orcohclix  (ilpi)i(i  (Elrod,  1901)  (Gastropoda:  Puhnonata: 
Oreohelicidae) 


H.  Lee  Fairbanks 

Pcnn  State  L'ni\ersit\' 
100  l)niver.sit\  Dri\p 
MdiKK-a,  PA  15001   I'SA 
lill  lCap.su.edu 


ABSTRACT 

rlic  icpii)ilucti\i'  aiialiiiiix  (it  spcciiiii-iis  cil  (h'iiiliili\  iiliiiiiii 
Iroiii  the  t^pe  loeaiitx  in  llic  Mi.s.siciii  Mountains  ol  western 
Montana  is  fij;in"ed  ani-l  eonipaied  with  that  ol  speeiiin'us  col- 
lected in  the  Swan  Mountains,  approxiniately  20  kilometers  to 
the  east.  It  appears  that  O.  ctlpina  is  found  in  hotli  the  Mission 
Mountains  and  the  Swan  Mountains,  which  ecmstitutes  a  range 
extension  lor  the  species.  In  addition,  the  reproductive  anato- 
niv  ofO.  (ilpiiiii  is  compared  with  that  olO,  suhnidis  iipaiium. 
Orcoliclix  (ilpiiia  lias  been  incorrectly  placed  in  Pilslins  () 
.suhnidis  liroup  ol  species  and  should  he  included  in  the  O. 
4-/rii^().v«  <j;roup 


INTHOUUCTION 

Tlie  lieiius  ()ri-()licli\  (Pilslm.  UJ()4)  is  cduiposcd  cil  a 
larije  iniinlu'r  ol  s]iccies  loniid  throimlioiit  tlu'  Hock\" 
Mountains  Ironi  southern  ("anada  to  soutlieru  Arizona 
and  Xcw  .Mexico.  .\  hin^v  anionul  of  iutraspecific  \aria- 
tioii  in  shell  shapt'  an<!  color  is  present  in  tlie  i;;('nus,  and 
it  is  olten  dilficult  lo  dclevniiiie  wlicllicr  the  eairse  of 
the  \ariation  is  cnvironniental  or  nenetie  (Filslnx.  1939). 
There  is,  however,  little  xariation  in  re|>rodneti\e  anat- 
oiTiy:  Piishrs-  (1939l  round  no  iclexanl  inleispeeilie  var- 
iation in  the  niorphologv  ol  the  nproduetixe  s\st(  in 
Nonetheless,  that  author,  based  in  pari  upon  llie  ralio 
ol  tlie  plicate  (ridijed)  portion  to  the  pusliilose  portion 
ol  tlie  internal  surlaee  of  the  penis,  allocated  s|)ecies  of 
Orcolirlix  into  one  of  three  groups,  llie  ()  s/r/ijosY/ 
group,  tlie  O.    i:hn((lis  group,  or  the  ()   i/iniipal  group. 

Elrod  (1901  i  d.scnlied  Orro/zc/a'  alpiiia  from  (lie  Mis- 
sion Mounlams  ■  .1  western  Montana  on  the  basis  of  the 
shell  only.  With  no  I  now  ledge  of  the  internal  aiiatoni\  of 
that  species,  l'ilsbi\  !,S9,)  tentatively  placed  O  iilpiiiii  in 
the  ().  siihniilis  group  becmse  O.  sithnidi.s  occurs  at  lower 
elevations  in  t]ie  saiai-  inountain  range  and  because  no 
members  ol  the  other  .'roups  occur  in  the  area. 

Speciuiens  of  Orcohclix  (dpiuc  from  the  txpe  localits 
had  not  been  collected  lor  neark  50  \-ears.  lieceiitK, 
however,  Hendricks  (199S^  collected  and  reporied  on 
several  specimens  from  the  t)pe  localit\,  some  of  which 


he  sent  to  rue  lor  dissection.  The  goals  ol  the  stuck'  were 

(1)  to  figure  the  reproductive  anatonn  ol  O,  olpiiid  and 

(2)  to  determine  its  taxonomic  position  relative  to  I'ils- 
bn's  three  Orcohclix  groups.  A  third  goal  was  to  com- 
pare additional  specimens  similar  to  O.  alpiiia  Irom  a 
localitx'  in  the  Swan  Mountains  to  those  Irom  the  t\pe 
localit\  ill  the  Mission  Mountains,  which  is  located  ap- 
pidxiniateK  20  kilometers  to  the  west. 

MATERIALS  AND  METHODS 

Specimens  ol  Orcohclix  iilpiiui  were  collei'ted  i Hen- 
dricks, 199S)  from  the  southeast  ritlge  of  St.  Maiys  Peak 
which  is  near  the  t\pe  localitx  on  the  southwest  ridge  of 
St.  Mans  Peak,  Tl'SN  H1S\\''S21  (Township,  Range,  and 
Section  Irom  USGS  topo  map).  Mission  Mountains, 
Pake  (JoinitN,  Montana,  elexation  approximatek  2S00 
meters,  on  27  Aug.  1997  (Figure  1 ).  .\dditioiial  speci- 
mens of  Orcohclix.  similar  to  O.  alpiiui.  were  collected 
b\  Bill  llamiiier.  on  7  -\ug.  1974  aboNC  Rumble  Lake. 
,Swaii  Mountains  {T2()N  RlfiW  S2),  Mis.soiila  Comitx, 
Montana,  elevation  approximatek'  2400  meters  (Figure 
1).  Two  topotxpes  and  three  siieeimeiis  from  the  Swan 
.Mount, liiis  vvei'c  dissected  .iiid  llieii  reproductive  svs- 
lenis  removed,  for  coniparalive  purposes,  the  repro- 
ductive s\  stems  ol  three  specimens  ol  ( ),  suhnidis  (ipdr- 
imii  Bern.  1919.  collected  on  2  Aug.  fy75,  at  Vellow 
Bav,  Flatiiead  Lake  (T24N  RI9W  S4).  Lake  CJoimtx, 
Montana,  elevation  ap[iroxiiu,itelv  S9()  meters),  were 
iisi'd.  The  reproductive  svstein  ol  each  specimen  was 
li.ieed  Irom  the  moiiitoi'  ol  a  dissecdiiLi  inieroscopeAid- 
eo  svstem,  llie  penis  ol  eaeli  speeiiuen  was  then  dis- 
seeled  lo  .illovv  loi  tracing  ol  theli  inlernal  snrlace.  Mea- 
surements ol  the  plicate  and  the  pusliilose  portions  ol 
(he  inlernal  peiiial  surlaee  wer<'  made  using  an  ocular 
iiiici'omeler.  llie  di.iwin^s  ol  one  s|')eciineii  were  used 
as  represeiilaliv<'  ol  llie  popiilalion,  I  he  diameler,  height 
ol  shell,  and  numlier  ol  vvlioiis  ol  the  speeinu'iis  Irom 
llie  Ivpe  loeaiilv  and  those  Irom  llie  Swan  Mdiinl.iins 
were  measured.  N'oiielier  specimens  ol  O.  ulpinii  Irom 
the  tvpe  Iocalit\'  and  from  thi'  Swan  .Mountains  are  de- 


H.  I..  F'airhanks,  2002 


Pasre  63 


1  MISSION  MTS 


Figure    1.     Wcvstiiii   Montana  sliimin'j;  tlic  Imatinns  nl   llic 
Mission  and  Swan  nionntains.  Stale  l)ar  =  2(1  km 


posited  ill  tin-  collt-ction  of  niollusks  at  tin-  National  Mu- 
seum of  Natural  lliston-,  Sniitlisoniau  lustitutidii.  W'asli- 


inatou,    D.C,    USA    (USNM    100233: 
1()02353). 


1.1  rsN\i 


RESULTS 

Siiolis  lidiii  till'  Mission  Moiilltains  ami  lioni  llic  Swan 
Mountains  arc  shown  in  Figures  2  ami  3;  their  inea- 
siiri'iuents  are  given  in  Tahle  1.  The  gross  reproclueti\t' 


%^  ^b 


Table  1.   Siicll  rncasnreincnts  lor  Orcolii'lix  alpiiui.  Measure- 
nienls  in  millimeters. 

IJkimelei   ol  \nnil)er  ol 

shell  Shell  heiijht  whorls 


Mission  Mountains 

Speeinien  A  8.0 

Specimen  B  7.9 

Specimen  C  7.5 

Specimen  U  8.4 

Mean  7.95 

Swan  .Mountains 

Specimen  .\  11,1 

Specimen  B  8.5 

Specimen  C  9.0 

Specimen  D  8.6 

Mean  9.:'5 


4.5 
4.6 

4.6 
4.57 


6.4 
5.0 
5.0 
5.2 
5.4 


3.3 
4.0 
3.5 
4.0 
3.70 


4.3 

4.25 

4.0 

4.0 

4.15 


*  Shell   was    [lamat^ed    dnriiiii   the   takint;   of  njeasurements. 
Hi'i'Jit  could  not  lie  olilaini'd. 


anatonix  III  individuals  troin  all  I  luce  localities  are  shown 
in  Figures  4-0,  and  the  inteiiial  penial  aiiatoiiiies  are 
shown  in  Figures  7-9.  Tahle  2  suniniarizes  the  iiiea- 
sureuients  ol  the  internal  penial  aiiatoun'.  The  plicate 
poi'tion  of  the  intt'rnal  penial  surl.iee  ol  the  Mission 
.Vhnnitains  (Figure  7i  and  the  Swan  Mnuntains  (Figure 
iS)  specimens  was  less  than  50',  (il  the  total  length  in 
contrast  to  that  nl  (I  suhriidi's  (iiuiriiiin  which  was  great- 
er than  oO'/r. 

DISCUSSION 

The  reprochietive  organs  of  Orcoliclix  alpina  from  tlie 
tvpe  locaiih'  (Figures  4  and  7)  are  t\]')ical  ol  the  genus, 
and  no  diagnostic  eharai'lers  at  the  specific  level  were 
detected. 

Pilshn'  (19.')0i  noted  thai  ni  the  ().  s/r/go.sY/  group  the 
plicate  poiiion  dI  the  p<'ins  was  ".  .  .  decidedlv  less  than 


Table  2.  Me.isiiremenis  ol  llie  plicate  porti<ins  and  imstnlose 
portions  ol  llic  mleni.il  smi.ices  ol  the  ])eni.scs  ol  Oivohclix 
tilpiiKi  .uid  ()   siilinidis  iijiiininii.  Measmemcnts  in  millimeters. 

Total         I'lisliilose        I'licate         Percent 
len^tli         ])ortioii         portion         jilicate 

O   (ilpiiia.  Mission  .Mountains 


Figures  2-3.  Shells  ol  Onolulix  ulpiiui  2.  From  the  tvpe 
localitv  in  the  Mission  Mninitanis  3.  Kiom  llie  Sw.ui  Moiin- 
tains.  Scale  har  ~  5  miii 


Speilnieii  .\ 

7..T 

4.1 

:5.4 

45 

Specimen  H 

10.0 

5.6 

4.4 

44 

( )   iiljiiiKi.  Swan 

Mountains 

Specimen    \ 

7.5 

4.4 

3. 1 

41 

Specmieii  B 

7.5 

4.6 

2.0 

.39 

Speeimeii  (  ' 

6.5 

4.0 

2.5 

38 

O.  sulinidis  (ipdiiinn 

HLF.59-1 

17 

7.5 

9.5 

56 

HLF.39-2 

15.5 

7.0 

8.5 

5.5 

11LF39-3 

14,5 

6,5 

S.u 

.5.7 

Pa"e  64 


THr:  NAUTILUS.  \o!.  llfi.  \().  2 


(iP 

F'igures  4-().  IvipiiKliu-tiM^  ssstcrns  ol  spctics  ol  Oirnlulix. 
4.  O.  (ilpitta  lioiii  llic  Mission  Mountains.  5.  (),  aljiiiiti  Ironi 
tlic  Swan  Mountains,  (j.  ().  siihriiilis  apdriiiiii  Iroui  tlif  Mi.s.sion 
Mts.  A(;  =  albumin  ^land;  K  =  epipliallns;  Gl'  =  genital  poR"; 
OV  =  ovotestis;  PH  =  [icnial  ivtiactor  muscle;  V  =  penis;  S 
=  spermatheea;  U  =  uterus.  Se;ile  liars;  l^'ignics  4  ;uul  5  =  1 
mm;  Figure  (i  =  5  mm. 


half..."  the  total  pcnial  Iciimli.  liiil  ^rcMlcr  ilian  luill 
tllc  total  peiiial  lciii;tli  in  hotli  the  ()  siihntilis  aronp 
and  the  O.  i/tiiajxii  groii[),  thi'se  two  gidiips  then  lieing 
separated  on  die  iia.sis  of  whether  the  ;inlerinr  hall  ol 
the  penis  \v;is  "  .  .  .  decidedly  .swollen     .  .'  or  not. 

Fiishn  originaliv  placed  O.  aljiiiid  in  the  ()   Mihnulis 
group  hecause  no  species  from  the  other  groups  (().  stri- 


Figures  7-9.  Internal  penial  analomx'  of  species  oi'  Orcolic- 
lix.  7.  ()  alpinu  from  the  .Mission  Mountains.  8.  O.  suhnulls 
(ipai-ium  from  tlie  .Mission  Mountains.  9.  O.  nlpinii  from  the 
Swan  .Mountains.  E  =  epipliallns;  C,V  =  genital  pore;  PH  = 
penial  retractor  mnscle;  P  =  penis;  VL  =  plicate  surface;  PS 
=  pnstulosc  surface:  \'  =  vagina.  Scale  bars  =  1  mm. 


H.  L.  F'airliaiiks.  2002 


Faire  fio 


oosa  and  O.  i/avnpai)  are  fbiiiul  in  tlic  Missinn  Moun- 
tains. Ho\ve\er,  tlic  lensrtli  of  tlie  jilicate  pciition  ol  tlic 
penis  of  ().  iilpuui  was  less  than  1/2  tot.il  [leiiis  length. 
whereas  tlie  [iliivite  poitioii  ol  (.).  siihnuli.s  (ijKiiiiiin.  also 
from  the  Mission  \h)untains.  was  <j;reatei'  than  1/2  total 
pi'nial  lentjth  (Table  2).  In  the  O.  tjUKquii  <j;r()n])  tlie 
plicate  portion  ol  the  penis  is  also  greater  than  1/2  total 
penial  length,  lint  in  addition  the  pro.xinial  portion  of  tin' 
penis  is  enlargeil.  which  is  not  the  case  for  O.  alpiiia 
(Figinv  4).  Therefore,  if  the  groupins;  s\'steni  of  Pilsiin' 
(19.39)  is  to  lie  lollowfd,  I  suggest  plai'emi'nt  of  (),  al- 
pina  in  the  (),  .sV/'igosy/  group. 

The  shells  ol  Orcniiclix  alpiiKi  lioni  the  .Mission 
Mountains  anti  the  shells  Ironi  the  Swan  .Mountains 
(Figiu'e  1)  could  not  he  separated  niorphologicalK  (Fig- 
ures 2  and  .3).  C.oinparisons  of  reproducti\e  svsteni  mea- 
surements for  these  two  populations  (Table  2)  did  not 
appear  signifieantK  different.  Therefore,  it  appears  that 


these  two  populations  belong  to  the  same  species,  and 
tliis  represents  a  range  extension  lor  ().  tilpiiici. 


LITERATURE  CITi:!) 

licrn.  .S.  S.  19H).  \Ii)lliisca  ol Clacicr  National  Park.  .Montana. 
Procccilint^s  ol  the  .Vcailciiiv  of  Natural  Sciences  ol  Phil- 
adelphia 71:  19.5-21)9. 

Elrod,  M.  |.  1901.  Montana  slulls.  Ho(k\  .Moiuilaiii  Ma<;a/.iiie 
2:  691-697 

Hendricks.  P.  199S  Krilisco\crv  ol  Discua  hndii.stini  Bern', 
19.5.5  and  OiciiIkIix  alpiiui  (Elrod,  1901)  In  the  .Vllsslon 
Mountains,  Montana,  with  comments  on  Orcniiclix  cirotii 
(PIlshiT.  191)01.  The  Nautilus  112:  .5S-62. 

Pllshn,  H.  A.  191)4.  (icnenil  Notes.  The  Nautilus  17:  1.31. 

Pllshr),  H.  A.  1939.  Laud  .Mollusca  of  North  .America  (North 
of  Mexico),  N'olunie  1.  I\u-t  1.  The  .-Vcadcuu'  of  Natural 
Sciences  ol  Philadelphia.  .Monographs  Niuuhcr  3,  Phila- 
delphia, wli  +  573  -I-  l\  IItuIcnI  pp. 


THE  XAITII.US  116(2):6r>-ft7.  2(){)2 


Pasie  66 


Notes 


Authorship  and  date  of  pubHcation  of  Ostrea  chilensis  Philippi 
in  Kiister,  1844  (BivaKda:  Ostreidae) 

Bruce  A.  Marshall 

Museum  of  New  Zealand  Tc  Pupa 

Tongarewa 
P().  Box"467 
Wellington 
NEW  ZEALAND 
BruceM@tepapa.go\-t.nz 


Ostrea  ch'dcnsis.  a  comnierfial  cnster  of  New  Zealand 
and  (>'liile  iBuroker  ci  a].,  1983;  Chanle\-  and  Dinamani. 
1980;  Jeffs  and  Creese,  1996;  6  Foighil  et  al.,  1999;  6 
Foighi!  and  Ta\lor.  2000),  eonsistentK  has  been  dated 
from  1845  nnder  anthorship  of  Philippi  in  all  references 
seen.  The  few  autliors  wlio  cite  the  original  reference 
for  O.  chilensis  (Dall,  1909;  Hanley,  1856;  Lamy.  1929) 
refer  to  Kiister's  edition  of  Si/stematisches  Conchi/Iien- 
Cahincl  ion  Maiiini  unci  Chemnitz,  p.  74.  pi.  13,  fig.  7, 
8.  in  N'ohniie  7.  part  1.  The  text  describing  O.  chilensis 
("Philipjii  in  Lit.")  is  part  of  Lieferung  186,  issueil  in 
1868  (Kiister.  1868)  (John.son,  1968:  366;  Smith  and 
Englami.  1937:  97;  Welter-Schultes.  1999:  185).  The  tax- 
on  Ostrea  chilensis  (as  "O.  [strea]  chilensis  Philippi")  in 
fact  originates  from  the  list  ol  species  depicted  on  the 
plates  inclnded  in  Lieferung  45  printed  on  the  wrapper 
thereof  (Kiister.  1844).  which  bears  the  imprinted  date 
1844  and  is  assumed  to  ha\(^  been  issued  then  in  the 
absence  olCxidence  to  the  contnuA  i  K]ZN.  1999:  Article 
21.3).  Here  Ostrea  chilensis  .st;intls  technically  as  a  no- 
men  nudum,  for  at  the  foot  of  the  list  is  the  statement 
"W'egen  Krankheit  des  Kuplcrstechers  k;um  Taf.  ,N111 
erst  niit  der  niiciisten  Lielernug  ;insgegebeu  wcrden" 
("Because  of  tlie  engraver's  illness,  Plate  1)  will  !)e  dis- 
triliuted  with  the  next  delixcn-.")  In  the  iibsencc  olexi- 
dence  to  tiie  contran  it  is  ;ippropii:ilc  to  :iccept  dial 
plate  13  was  indeed  issued  in  the  next  Lieferung  (which 
is  also  imprinted  1844  and  assumed  to  haxc  been  issued 
theni.  and  that  the  name  ().  chilensis  became  available 
fnjm  the  nomcnclatural  standjioiut  at  tlu>  time  of  issue 
ol  that  Licfenm,;.  I  Note  that  pi.  13  was  erroneonsK  at- 
tributed to  Lielciuug4l.  and  Lieferung  45  erroueonsK' 
dated  1843  by  Welter  S.hultes  ( 1999:  185),  but  the  latter 
correctly  dated  elsewliere  in  the  compilation.]  Although 
wrappers  for  this  work  were  intended  to  be  discarded, 
and  most  indeed  were,  tax;i  introduced  on  ihem  with 
reference  to  illustrations  (uKill  criteria  for  availabilit\- 
(ICZN  Articles  8.1,  12).  Tivxa  introduced  on  these  wrap- 
pers were  acceptetl  as  published  b\  C.  D.  Sherborn  (in 
Smith  and  Engkind,  1937:  89.  '  nomen  el  /igifw").  who 
failed,  howexer,  to  include  Ostrea  chilensis  in  Index  An- 


iinaliuin  (Sherborn.  1922-1933).  The  authorship  and 
date  of  publication  of  O.  chdensis  are  thus  Philippi  in 
Kiister,  1844  (Kiister,  1844). 


.ACKNOW  LEDC;-\1ENTS 

For  checking  accession  records  tor  parts  of  Si/steina- 
tisclies  Conchi/Uen-Cabinet  von  Moiiini  und  Chemnitz 
and  other  works.  I  am  grateful  to  R.  Janssen  (Sencken- 
berg  Museum,  Frankfurt).  .-K.L.  Kiibat  (Washington, 
D.C.),  D.G.  Reid  (The  Nattual  Histon-  Museum,  Lon- 
don), A.H.  Swann  (Wheldon  and  \\esle\-  Ltd.,  Leighton 
Buzzard),  A..  Waren  (Swedish  Museum  of  Natunil  His- 
ton-. Stockholm),  and  F.  W.  \\'elter-Schultes  (Unixersitiit 
Gotiugen).  Special  thanks  to  S.  Jordan  (La  Habra 
Heights,  California),  who  supplied  a  photocopv  of  a 
wrapper  of  Lieferung  45,  and  to  P.  Bouchet  (Mnsenm 
Niitional  ilHistoire  Naturelle,  Paris)  and  A.  L.  Kabat  for 
extensive  comments  on  the  subject. 

LlTER.JiTURF  CITED 

iiiu.ikcr.  N  K,.  W  Chanlev,  If  |  Cnmlidd,  1",  Diiuini.ini.  19S.3. 
Svsteniatic  status  of  hvo  instcr  populations  oi  the  ijenns 
Tiosirrd  honi  New  Ze;ilancl  and  Ohilc.  Marine  Biologv  77: 
l()l-:2()(), 

C.'lianliv.  V.  and  T.  Dinamani.  19S().  ( .'onqianitive  descriptions 
oi  some  ovster  lanae  irom  New  Zealand  and  (,'liile.  and 
a  description  ol  ;i  new  i^cnus  ol  ovster,  Tiostrcii  New  Zea- 
land |onrnal  ol  \1. nine' and  Freshwater  Heseauli  It:  103- 
120  ■ 

D.ill,  W  II  U)09.  Report  on  :i  collection  of  shells  Irom  I'c'ni, 
Willi  .1  snmniai'V  ol  the  littoral  marine  Molliise:i  ol  the 
l'rriivi:m  /oologieal  province.  I'loceedings  ol  th<'  I'nited 
SI. lies  National  .Museum  .')7:   147-294. 

Ihiiilc-v.  S.  lS12-lS.5fi.  .\n  illnstiMlcd  ,ind  dcsiriplive  citaloune 
ol  Kecc'iit  bivalve  shells  Willi, mis  and  Nor<;:ite.  London. 
W'iii  +  .'W2  pp. 

UrZN  (Inteni.itional  ('omimssioii  on  Zooloijical  Xonniiilie 
lure).  1999.  Inteinalioiuil  (lode  ol  Zoolos^ical  Nomencla- 
ture. Fourth  Edition.  International  Trust  lor  Zoological 
Nomcnrhiliirc.  London.  .306  pp. 

Jells,  \.  C.  and  IL  C.  Ca'cese.  1996.  Ovcimcw  and  hihliogra- 


p.  Cal 


ZOO 


'd''v  67 


pli\  (it  research  on  the  Chilean  cnstcr  Titistrcd  rhilviisis 
il'hilippi,  1S45)  from  New  Zealand  waters.  |i)urnal  ol 
Shellfish  Research  15:  305-311. 

|iiliiiMiii.  H.  I.  196(S.  Martini  and  ( 'heninit/ iKiKslers  cilitionl 
S\steniatisches  Conchvlien-Calmiet.  1S37-192().  a  com- 
plete collation.  Jonmal  ot  the  Societ\  lor  the  Bil)lio^r.ipli\ 
of  Natnral  Histiin-  4:  363-367. 

Kiistcr.  H.  C.  1844.  Inhalt  [list  ot  species  depicted  on  plates 
inclnded  in  Liefennig  45:  pi.  13,  howexcr,  exidi'utK  issned 
ni  l.icrcrun'.i  46.  1844].  S\steniatisches  (  jiiicli\  lii'ii-(  .'ahi- 
nrt  \oH  .Martini  nnd  Chenniitz  (Kiister,  II. (^  cd.^  7  i  I  )  45 
wrapper  Baner  and  Raspe.  Niirnherg.  (Phot(K-op\  in  Mol- 
liisca  Section  libran'.  Mnsenni  ot  New  Zealand  Te  I'apa 
Tonsjarewa). 

Kiister,  H.  C.  1868.  Conchifera.  Zweischaline  Mollnskt'ir  ( 'mi- 
chitera  inononi\aria.  .Mnschelfhieri'  niit  eineni  Sclilii'ss- 
nniskeln.  Svsteniatisches  C'oncli\lien-CJabinit  \i>ii  Martini 
nnd  Chenniitz  (Kiister.  H.C.  ed.)  7(1)  186:  5(1-S4.  Baner 
anil  Raspe.  Niiniberg. 

Lani\.  Iv   U)29.  Revision  des  Ostna  \i\;tnts  dn  Mnsruni  N:i- 


tional  d'llistoirc  Natnnllc  dv  Paris.  |oin'nal  de  (.'onch\- 

liolot^ie  73;   1— 4(). 
6  Foighil.  I)  .  B,  .\.  .Marshall,  i    j,   llill.ish  and  M.  .\.  I'ino. 

1999,  Transd'acihe  ranije  c\ti-nsion  1)\  raltini;  is  iiilerred 

for  the  Hat  osster  ()\lrici  c/ii/r/is/s.  IMologiciil  Bnllctin  196: 

122-126. 
(J    Foighil,    I).   :Liid    I).    |.    I'.ivlor    2lll)(l.    I'Aoliitioii   ol    parent. il 

care  and  oxnlation  lielia\i(>r  in  ovslcrs.  .Molei  iifii'  l'li\lo- 

genetics  and  K\ohition  15:  .301-313. 
.Sheriiorn.   C     O     1922-1933.    Inde.\   .-Vninialinni    1801    IS50, 

British   Miiscuni  (Natnral   lliston),   London;  Longmans, 

Green  ami  Co..   B.   OM;iritch.   Dnlan,  ()\lord  l'ni\crsit\' 

Press.  Wheklon  and  Weslew  and  Oliver  and  Bo\il. 
Smith,  E.  ,\,  and  11.  W.  Kngland.  1937.  NLulini  and  Chemnit/ 

( tinester's    edition'    Sxstematisches    Conch\lien-Cal)inet. 

1837-1918.  Jonrnal  of  the  Si)ciet\  lor  the  Bililionraphx  of 

N:itin:il  IlistoiA    1:  89-99. 
Wclter-Sdiiillcs    I"  \\     1999.  S\ stenKitisches  Conch\lien-Cal)- 

inct  \on  M.iitmi  mid  ( 'licnmit/  i  1  S37-I920'.  !iil)liograph\ 

of  the  Milnmcs  m  (Kiltingin    Archives  ol  N:itnral  lliston 

26:  157-21)3. 


Tadashicre  Habe  (1916-2001 


Paul  Callonion 

Department  ol  Malacolo'j;\ 
The  .\cadem\  ol  Natmal  Sciences 
Philadelphia.' P,\  19103-1195  US,\ 
callomon(£!'acnatsci.org 


Tadashige  Halie,  who  died  at  S5  on  Dccciiilier  29.  2001, 
was  one  of  the  most  prodnctixe  and  iiilhieiitial  /odlogists 
of  the  twentieth  tentun'.  He  plaxed  a  nia|o|-  pait  in  tlie 
development  of  malacologA'  in  [apan  anil  the  western 
Pacific,  and  authored  a  large  mimlier  ol  hooks  that  set 
the  stanikuiis  \nv  the  stniK  ol  the  swstematies  and  dis- 
trihiition  ol  niciliiisks  Ironi  that  area  ol  the  wcirlil, 

l)i.  lialie  was  horn  March  31.  191fi,  in  the  village  of 
Hioki,  part  of  what  is  now  Sasavama  in  Hyogo  Prefec- 
tnre.  He  attended  Ikeda  High  School  in  neighhoring 
Osaka  Prefecture,  sta\ing  on  to  wmk  as  an  assistant 
teaclier  until  1939,  Electing  to  stiidv  zoologx  at  Kvoto 
Imperial  Universit^■  from  1939  to  1941,  he  then  trans- 
ferred to  Seto  .Marine  Lahoraton  in  Sliiiali;inia.  return- 
ing to  Kvoto  in  1945.  Dr.  Hahe  had  joined  the  stall  :it 
K\'oto  as  a  junior  researcli  assistant  in  1942  and  liecanie 
a  full  research  assfstant  in  1949.  From  194fi  to  1950  he 
also  served  as  an  adviser  to  the  Natural  Resources  .Sec- 
tion ol  .\llied  (U'lieral  Heaili|uarters  in  Tokvo. 

.\lter  receiving  his  doctor  ol  science  degree  ni  195i. 
Dr.  Haiie  was  appointed  assistant  professor  at  Kviisini 
Universit\'  and  moved  to  the  Maiine  Biological  Laho- 
raton' on  .Amakusa  Isl.ind  in  westi'rn  Kvushu,  lie  re- 
tained his  position  at  Kviishn  liniversitv  lor  five  vears 
after  moving  ni  1962  to  the  National  Science  Museum 
in  Tokvo  (NSMT). 


Dr.  Hahes  career  at  tlie  NS.MT  lasted  from  his  ap- 
pointment as  researcher  in  1962  to  his  retirement  as 
head  of  the /oologv  depiirtnient  m  19S0.  I  le  then  seneil 
for  four  vears  as  a  professor  at  the  Ocean  Institute  ol 
Tokai  Liniversitv.  where  he  remained  |irofessor  emeritus 
until  19S9.  From  1985  to  19SS  he  w:is  also  director  of 
tlie  Tokai  L'niveisitv  Museum  of  N.itnial  llistor\.  In 
19S6  Dr.  llahe  was  awanled  the  Order  of  the  Sacred 
Treasure  iZuilio-shoi  |,\  the  |apanese  government  lor 
his  contributions  to  /oological  reseaicii  in  Japan. 

Dr.  ilalie  lirst  puhlished  formally  as  co-author  ol  a 
1935  report  on  shell  collecting  in  .Manchuria  and  Korea. 
For  more  than  hftv  veais  he  wrote  |)roi!igiously.  alone 
.ind  in  cd-.inthorshiji  with  all  the  other  senior  Japani'se 
inakicologists  of  the  post-v\ai-  period.  Several  ol  iiis 
hooks  are  well  known  outside  japan,  including  ,S7(('//.s  of 
lite  Wcslcni  Facijic  in  Color  il964i  and  S/(c//.s  of  tlw 
World  in  Colour  (\ol.  1.  19(i5.  with  Kivoshi  Ito;  \ol.  2, 
1966,  with  Sadao  Kosnge), 

Pearlier  in  his  career,  Dr,  llahe  worked  closelv  with 
Toknliei  Kuroda  (  1SS6-  19S7  .  whii  had  arrived  at  Seto 
M;Mine  Lahoralorv  in  1940  and  was  to  have  a  formative 
inllnenee  on  tlie  vounger  man.  The  i\\o  published  their 
first  joint  book,  a  svnopsis  of  the  Japanese  Helicac(>a,  in 
1949,  This  woik  was  followed  In  the  exten.sive  Illustrat- 
ed Catidoiiuc  of  }a])(iiu'sv  Shells  (1949-1955)  and  the 


I'.iilL'  bb 


TIIK  NAUTILUS,  \ol.  IKS.  No.  2 


landiiiark  Checklist  find  Bihlionriipluj  of  the  Remit  Ma- 
rine Molliisca  of  Japan  i  1952).  Tlic  latter  was  piililishetl 
ill  part  as  a  result  ol  its  Dr.  llalie's  work  tor  .Vllii'd  Gen- 
eral IIeacl(|uarters.  and  pro\ided  the  first  thorough  col- 
lation oi Western  and  local  literature  on  Japanese  Mol- 
lusca.  In  1971  Kuroda  and  Habe  were  joined  In  Katsura 
0\ania  (1917-1995)  in  producing  the  lavish  The  Sea 
Shells  of  Sa'fiami  Bat/.  This  extensive  work  owed  its  ex- 
istence in  part  to  the  favorable  relationships  all  three 
atithors  enjoved  with  the  Imperial  household,  a  connec- 
tion further  reflected  in  a  number  o(  subsequent  papers 
lo-authored  b\  Dr.  Habe  and  Prince  Hitachi  (Masahito), 
brotluM'  of  the  current  emperor. 

During  the  four  decades  that  followed  tiie  .Second 
World  War,  [apanese  malacologv  was  to  a  considerabk' 
extent  sustained  by  the  sponsorship  antl  participation  of 
a  inunber  of  keen  amateur  collectors  and  authors.  Dr. 
Habe  was  an  enthusiastic  advocate  of  collaboration  with 
amateur  collectors,  and  skillfulK  helped  chamiel  the  en- 
ergv  and  resources  of  benefactors  such  as  Tetsuaki  Kira 
and  Rvosnke  Kawamura  into  tlie  production  of  useful 
publications  and  scientificalK'  significant  collections.  He 
was  instrumental  in  obtaining  the  vast  Kawaunu'a  collec- 
tion lor  the  NSMT,  and  co-authored  an  impressive  iio- 
nographv  based  on  it.  World  Seashelh  of  Raritij  and 
Beantij  (1991). 

Dr.  Habes  specialitv  was  taxonomy,  and  he  was  the 
first  Japanese  malacologist  to  sene  on  the  council  of  the 
Internationd  Trust  for  Zoological  Nomenclature.  In 
1977.  he  published  Si/stcmafics  of  Mollusea  in  japan: 
Bivalvia  and  Seapliopoda.  a  complete  iliustratetl  taxo- 
noniical  treatment  of  both  groups. 

.Another  important  but  less  well-known  lacet  of  Dr. 
Habes  career  was  his  work  in  translating  and  sunnnariz- 
ing  developments  in  Western  malacologv  for  the  benefit 
ol  his  Japanese  colleagues.  This  took  the  form  of  a  long 
series  of  well-researched  biograjihies  of  Western  mala- 
cologists  and  zoologists  as  well  as  reviews  of  books  and 
monographs  aTuI  rejiorts  on  Japanese  l\pc  spi^cimens  in 
overseas  nmsenni  collections. 


He  was  v\'ell  known  among  malacologists  in  the  West, 
and  traveled  overseas  more  olten  tlian  mauv  of  his  con- 
ti'inporaries.  During  his  peiiod  with  Allied  (General 
Headquarters,  he  worked  both  with  Dr.  .\lv-in  ("ahn  and 
Dr.  Mvra  Kec'u  of  Stanford  Universitv.  In  later  years,  he 
was  a  ri'gulai'  visitor  to  the  United  States  and  a  frienii 
in  particular  of  Di'.  H.  Tucker  Abbott,  with  whom  he 
shared  a  broad  outlook  concerning  amateur  scientists 
and  popular  publications.  \'isitors  to  the  NSMT  during 
Dr.  Habes  tenure  alv\avs  bi-nefited  troni  his  renowned 
hospitalitv. 

A  member  from  its  early  davs  of  the  Malacological 
Societv  of  [apan.  Dr.  Habe  served  as  vice  president  from 
I9fi:3  to  1979,  as  president  from  1979  to  1995  and  as 
emeritus  president  until  his  death.  In  recent  vcars  his 
failing  evesight  began  to  restrict  his  activities,  but  lie 
remaineil  an  active  contributor  to  conlerences  and  pe- 
riodicals until  the  lati'  199()s. 

In  the  course  of  his  career.  Dr.  Habe  introehiced  more 
than  a  thousand  new  names  to  science.  Collations  of  his 
new  taxa  wt-re  published  bv  Inaba  and  Ovarna  (1977) 
and  Okamoto  (2001).  The  latter  work  is  complete,  in- 
cludes a  full  bibliograpln  and  cites  976  new  species 
names  anil  .'ilo  new  genera.  .-\  partial  bibliographv  was 
publishedlnDr  llal>e  himself  in  19S(). 

IJTKH.VniRE  CITED 

llahc.  T,  t9S().  Bibliographv  of  Tadashigc  Halic.  T  llahe.  To- 
k-vo,  47  pp.  [Partlv  in  Japanese.  | 

Inaba.  T.  and  K.  Ovania  (Eds.)  1977.  ( Catalogue  ol  nmlliiscaii 
ta\u  described  In  Tadasliige  Hahe  during  1939-1975. 
vritli  illustrations  ol  hitherto  uiifi<;nre(l  species.  Okiiiaehi- 
su-k;u.  Chiba,  185  pp..  7  jils. 

Okamoto,  M.  (ed.)  2001,  .Molliiscan  ta\a  tlcsinhetl  li\  Tada- 
shige  Hahe.  \'i>l.  1.  (Gastropoda  (excluding  lleterohran- 
chia):  i-wiii  +  1-S91.  \'ol.  2,  (lastropocki  (Heterohran- 
chia).  Cephalopoda,  Bivalvia  &  Scapliopo(hi):  892-1630  -1- 
xix-vxxix  +  index  1—42  +  xl-xli.  Connnittee  lor  celehrat- 
itig  Dr,  T.  Habes  eightieth  birthdav,  Tokvo.  [Partly  in  Jap- 
anese, I 


THE  XALTIIA'S  1  lfi(2):fi9-7().  2002 


Paa-  69 


Book  Review 


Panamic  Proiiuce  Molhiscdti  Literature: 
Additions  and  Chauges  from  1971  through 
2001.  111.  Gastropoda. 

Sko^liind.  C\  2002.  Fananiic  Pnnincf  Molliiscaii  Littr- 
attire:  Aiklitioiis  and  C'liautics  from  1971  tlin)ii!j;ii  2001. 
III.  Gastropoda.  Tlie  Festi\ii.s  33  i  snpplcinent):  i-\i  + 
1-286  pp. 

PossibK'  inspired  h\  the  ininiensits'  oi  A.  M\Ta  Keen's 
Sea  Shells  of  Tropical  West  Aiiicrica  (Keen.  1971)  and 
elearK'  based  on  a  liietime  of  stncK  ol  tlie  subject,  lor 
\ears  C^arol  Skoiilnnd  has  been  updatin'j;  the  in\enton' 
of  Panamic  niollnsks  as  ori<j;inalK  jiublished  in  Keen's 
wiirk. 

In  the  couchision  ol  a  recent  ie\ie\\  ol  Skoglunds  \ol- 
unie  on  bivaKes  and  chitons  I  Leal,  2001,  re\ie\\' ol  Sko'j;- 
knid,  2001  >'  1  had  wished  that  the  autlior  eonld  provide 
an  equixalent  npilate  on  tlie  gastropods.  That  work,  also 
pnblished  as  a  supplement  to  The  Fi\linis_  is  now  avail- 
able. The  exhaustive  compilation  results  from  vears  ol 
record  keeping,  as  attested  to  b\  the  long  list  ol  perti- 
nent publicati<ins  In  the  author  in  that  periochcal.  autl 
SkogluiKl's  nnqnestionable  grasp  ol  the  subject  of  Pan- 
amic mollusks.  The  general  lorniat  follows  tint  adopli'<l 
for  the  previous  parts.  Although  most  ol  the  updates 
consists  of  the  inclusion  of  new  species.  Skoglnnd  also 
presents  newlv  adopted  use  of  svnonvnis,  new  combi- 
nations (these  latter  t^vo  categories  indicated  bv  Keens 
original  species  number),  as  well  as  thi'  occurrence  of 
supraspecific  taxa  new  tor  the  region  (indicated  in  bold- 
lace). 

In  a  similar  lashion  to  die  volnme  on  bivalves  and 
chitons,  the  new  publication  is  spiral-bonntl  and  [iro- 
tected  bv  a  plastic  cover  Although  the  compilation  is  not 
illustrated,  reference  to  original  works  and  existing  fiti- 
ures  is  abrmdant  and  seems  to  be  complete. 

Like  the  previous  volume,  the  gastropod  update  cov- 
ers the  coast,  shell,  and  slope  between  25°  N  and  fi°  S. 
Most  welcome  bonuses  are  the  extensive  treatments  of 
mollusks  from  oceanic  islands  in  the  tropical  easterii  Pa- 
cific, (e.g.,  Clipperton.  Cocos,  and  Ciorgona)  and  ol  those 
associated  with  geologicallv  active  features,  sucli  as  hv- 
drothermal  vents  and  cold  seeps,  of  the  adjacent  deep- 
sea  areas.  The  onlv  oversight  I  noticed  in  a  brief  rea(hng 
of  the  suprageneric  taxa  were  tiie  misspellings  ol  "(Joc- 
cnlilormia"  [Cocculiniformia]  (pages  ii,  23i,  as  a  subor- 
der under  the  subclass  Neritimoipha.  Skoglnnd  used  the 

'  luadverteiitlv,  in  the  title  ol  iiiv  review  I  citeil  the  liivalve  anil 
chiton  volume  as  ",  .  .from  197i  to  2(1(11  vvIkmi  it  slioiild  have 
been  ".  .  .from  1971  to  2()(»(l,"  M\  apoloLjii's  to  the  autlior  , mil 
editor. 


correct  s[)elhng  for  the  subclass  (^occnliniiormia  (same 
pages),  and  1  wonder  whether  the  misspi'lli-d  woril  was 
meant  to  be  "( .'oci  iiliiioida  oi"  a  re|)etition  ol  "('occii- 
liniformia.  " 

The  hierarchical  lanked  anangeiiicnl  ol  lainilies  and 
genera  generallv  lollows  that  adopted  bv  Keen.  In  manv 
cases,  however,  those  were  modified  through  the  atlop- 
tion  of  more  recent  taxonomic  rearrangements,  inelnd- 
ing  some  ol  the  groupings  nltimatelv  compiled  b\' 
N'anght  (19SS)  and  the  inclusion  of  additional,  mostiv 
deep-sea,  lainilies  anil  geiier.i  Mi.il  h.ive  been  intro- 
duced in  the  last  30  vcirs.  iiiainlv  through  the  ellorts 
ol  |.  11,  Met, can  anil  others.  'I'lie  ailvaiK'cs  in  our  nn- 
derstaiRling  ol  gastropod  phvlogenv  over  the  past  lew 
decades  (e.g..  Ponder  and  Lindberg,  1996.  1997)  have 
resulted  in  a  higher  classification  of  the  Class  Gastro- 
poda that  differs  significantiv  bom  that  incoiporated  in 
earlier  works.  While  this  new  classification  is  being 
adopted  in  manv  recent  works  on  gastropods  (e.g., 
Beeslev  et  ak.  1998;  Higo  et  ak,  1999;  Okutani,  2()()()) 
the  opportiinitv  to  update  the  taxonomic  arrangement 
of  Panamic  gastropoils  has  been  missed  in  Skoghmd's 
work.  Tliis.  however,  shall  not  be  a  major  handicap;  tiie 
author's  chief  objective  is  to  give  a  consistent  update 
on  the  inventon  and  nomenclature  of  Panamic  gastro- 
pods, providing  links  both  to  Keen's  book  and  the  orig- 
inal references,  lii  mv  opinion,  she  has  achieved  what 
she  set  out  to  do  with  great  success.  If  voii  use  Mvra 
Keen's  work,  voii  must  have  these  updates. 

The  cost  in  the  US  is  S35  postpaid,  overseas  S5()  (air- 
mail onlv  1.  Contact  the  San  l)ie',io  Shell  Club.  %  3883 
Mt.  Blackburn  .Ave..  San  Diego,  CA  92111,  USA,  or  e- 
mail  Carole  Hertz  (cmhertzC«pacbell,nel)  for  further  in- 
lorm.ition. 

LITERATUHK  CATED 

Beeslev.  P.  1...  C.  |.  ti  Ross  and  A,  Wells  <e(ls.  i  199S.  Molliisca: 
the  Soiilhern  Svnihesis.  Fauna  of  .\iistralia.  \'oI.  5.  CSIKO 
Puhlislmii;.  .Melliourne.  .5fi'5  pp.  Part  \l  pp.  565-1234 
pp  (Part  B' 

Higo.  S..  P  Callomon  and  V.  (ioto.  1999.  Catalogue  and  Bih- 
hographv  of  the  Marine  Shell-hearing  Moliusca  of  Japan. 
F.lie  Scicniihc  I'nlilications.  Osaka.  749  pp. 

Keen.  A.  \1  I97F  Sea  Shells  of  Tropical  W'e.st  America:  Ma- 
rine Moflusks  from  Baja  California  to  Peni,  2'"'  edition. 
Stanford  L'niversitv  I'rcss,  Stanford,  \iv   +   1064  pp. 

Leal,  J,  II.  2001.  liook  review:  Panamic  Province  Molluscan 
Literature:  .\dditions  and  Chani;es  from  1971  throui;li 
2001  12000].  The  Nautilus  115:  .37. 

Okutani.  'I",  leifi  2000.  .Marine  Mollusks  in  Japan.  Tokai  Lni- 
versitv   Press,  Tokvo,  xlviii   +   1173  |-H2]  [)p. 

Ponder.  W.  F  .mil  1),  Lindberg.  1996.  Gastropod  phvloiienv: 
challenges  lor  the  9()s.  in:  J.  I).  Tiylor  (ed.)  Origin  and 
Evohitionai-v  K;idiation  of  tin-  Moliusca.  Oxford  University 
Press,  Oxford,  pp.  135-151 


Pagr  70  THE  NAUTILUS.  \ol    1  Ifi,  No.  2 

Ponder,  W.  F.  and  D.  Lindbcrg.  1997.  Towards  a  plixlogcnvol  N'anglit,  K.  (,'.   1988.  .\  (;Ia.ssification  of  tlic  U\intT  Mollnsca. 
gastropod  niollu.scs:  an  anaKsis  using  niorpliological  cliar-  .Kniciican  Malafologists,  Inc.,  .Vlelbonnie  [Florida],  xii  + 

acters.   Zoological    |(iurnal   nl    llu-    Liiuican   Socii't\    119:  189  pp. 

•'^^-seo.  j^^^.  ^^  Leal 

Skoglund,  C.  2001.   Fanaiuic  l'i(.\MKv   \l.,llnscan   Literature:  -[-|,,.  Hail,.v-.\l:ittli,.\\s  Shell  .Musnnn 

Additions  and  (,'lianges  troni  1971  throngh  2000.  1.  Bi\-  p(^   gox  1.580 

alvia.  2.  PoKplacophora.  The  Fcstixiis  32  (supplement):  Sanibel,  FL  .>59.57  US.\ 

i-v  +   1-119  iBi\al\ia):  1-20  i  PoKplacopliora).  jleal@.slicllnins(inn.org 


This  piihlication  sponsored  in  part  1)\ 

till'  State  of  Florida.  Department  ol  Stale 

Division  ot  C^ultural  Affairs,  anil 

the  Florida  Arts  Couneil 


Tliis  issue  sponsored  in  pait  1)\: 


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TH  E€7NAUTI  LUS 


Voltimc  116.  Siimhcr  3 

Srpfcinhcr  30.  2002 

ISSX  002H-1344 


CONTENTS 


S\en  \.  Nielsen  Tertian  Xt'iioplidriilae  (Castropixla^  ol  western  South 

Tlionias  J.  DeN'nes  America 71 

Jose  VVillibaldo  Tlioine  Hedescriptioii  ol  tlie  (j;emis  and  species  llvtcrovainniiui 

Suzele  Kodrigues  Cioiiies  liiiuit/aua  (Lesson,  1830)  (Gastropoda:  Soleoliiera: 

Rosanc  Souza  da  Silva  Wronicellidae) 79 

Kristiina  0^"aska  Aiiatoiiix  of  the  droiiiedai-\   juinpin'^-shii^.  livinplidlUi 

Lvie  C^liichesler  ilromcdinius  Bi'aiisou,  1972  (CJastropoda;  St\loiiiiii:iliipliora: 

Heike  Reise  Arionidae),  with  new  distributional  records S9 

William  P.  Leonard 
Jim  Haui^h 

Philippe  Boiiehet  New  species  ot  deep-water  Caneelhiriidae  ( (laslropoda^ 

Richard  Petit  from  the  soutliwestern  Pacific 95 

Guido  Pastorino  Spawn  ol'  the  Pata^oniaii  nastropotl  I'liriiilliria  phunhcd 

Pable  E.  Penchaszadeh  (Pliilippi,  1S44)  (Bnccinidae) 105 


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THE  NAUTILUS  116(3):71-7.S,  2002 


Paije  71 


Tertiaiy  Xenophoridae  (Gastropoda)  of  western  South  America 


Sven  N.  Nielsen 

(ii'ciIdt^iscli-Palaontiilot^isrlics  liislitiil 

and  Must'iini 
Universitat  Hanilmrg 
Bundesstrasse  55 
20146  Haiiibuis; 
GEKMANV 
iiielsenS'geowiss.iiin-luiiiihiin^.tlc 


Tlioinas  J.  I3e\'ric's 

liiirkc  Miisiiiiii  III    \aliiial   llislon   .iiid 

Ciilliiiv 
I  ni\ciMl\  (il  Wasliiii'^tdii. 
Sraltic,  W  \  9.SU)5  USA' 


Al^STHACT 

Tlii'cf  spcrifs  ol  Xi'iiDplimular  aic  ri'|)(irlc'il  Imiii  die  TcrtLiiA 
ol  western  Soutli  Amenca:  Xiiiopliord  iiiniitiiS.i'rii  new  species 
from  the  Oligoeeiie  of  Peru.  Xciiophnni  piniliiiHf  new  species 
from  the  Miocene  Na\idad  Formation  of  central  Chile,  and 
Stclliiiifi  kririicrhiiiihdhli  new  species  from  the  Tertiar\  ol  Pen- 
insnla  .\ianeo.  sonth-eential  (  'hile.  .\11  ol  tliese  species  are  first 
recortls  ul  .Xenophoiidae  lor  tlic  respe(.ti\e  conntrii's  'iTie  rc- 
lationsliip  ul  Sunlli  Aniernan  \i  iiiijilioni  species  wjlli  'I'ertian 
New  Zealand  tiL\a  and  tlie  tlillicult\  in  identilxTng  a  species 
closelv  related  to  the  other  .South  American  Siclltiria  are  dis- 


INTRODUCTION 

Xeiioplioridae  ar<'  a  u;iniip  i>l  cxclusivcK  wann-water  ma- 
rine gastropods.  The  liioloin,.  fossil  liiston.  and  taMiiioinv 
of  25  recent  species  has  heen  re\ie\\ctl  In  Ponder 
(19S3).  who  recognized  the  single  genus  Xnioplidrti  Fi- 
scher \()n  W'aldlieini,  1807,  including  the  subgenera 
Xciioplioni  sciisii  sfriiiii.  Oiiiisfn>;  Swainson,  ISdO.  and 
Stclldrid  Schiniclt.  1S:52.  IIowe\cr.  most  workeis  now 
recognize  these  at  the  generic  rank  (e.g..  Kicijil  and  AH, 
1999).  a  \iew  that  is  followed  liere.  Where  known,  the 
protoconc'h  in  Xcnoiilioni  species  consists  of  aliont  .IS 
low  trochospiral  whorls  I  Bandel.  199:3:  pi.  12.  Hg.  1),  a 
t\pe  of  moq^liologx  that  indicates  planktotropliic  (le\i-l- 
opment,  which  in  tiu'n  suggests  long-distanc'e  dispeisal. 
Ponder  (19S.3I  also  described  Tertian  fossil  species 
from  Australia,  wliile  in  an  earlier  work.  Ben  (1977)  re- 
viewed the  Onozoic  Xenophoridae  of  New  Zealand 
which  include  an  Eocene  to  Miocene  species,  a  Miocene 
species  and  a  Pliocene  to  Recent  species.  Until  now,  Ce- 
nozoic  Xenoplioridae  ha\e  never  been  recorded  from 
Pern  or  Chile. 


'  Mailniu  ackhess:  Box  13(161.  Burton.  WA  9Sl)f3  USA. 


(;k()1.()(;y  of  fos.sif-hfahixc  i,(K,Ai,rriFs 

Pi.sco  B,\si\,  Pkhh 

The  Pisco  f^.isin  of  southern  Pern  i  Fignri'  1 )  is  a  forearc 
basin  (Unnbar  et  ah.  1990)  with  four  tli'positional  se- 
quences containing  upper  Eocene.  Oligocene.  lower  to 
miildle  Miocene,  and  middle  Miocene  to  lower  Pliocene 
shell  and  littoral  deposits  i  DeWies.  1998).  Specimens  of 
Xiiiojihiiii:  are  most  often  found  in  ma,ssi\'e,  bioturbated, 
medinm-grained  sandstones  of  the  Oligocene  Otnnia 
Formation  behveen  Paracas  and  i5aln'a  de  la  Indepen- 
dencia  (Figure  1),  together  with  \aKcs  of  Canlilti  iicw- 
clli  Ri\era.  f957.  The  .\'r/i(i/)//(i/v/-bearing  sandstones  in- 
lerpicted  as  shallow -w  .tier  nearshore  slielf  deposits, 
overlie  a  basal  transgressi\f  si'i|nence  of  nu>dium-bed- 
ded  coarse-grained  sandstones  with  uearK  monospecific 
mollnsean  assemblages  of  rtirrilcllii  or  Ostrcd.  and  nn- 
ilerlie  a  thick  se(|ueuee  ol  thin-bedded,  tuffaceons.  fine- 
grained siltstones  with  thin-shelled  \al\<>s  ol  Chlmiujs 
and  fish  scales  of  sar(fiues  and  anchovies  (DeWies, 
I99S1. 

Internal  molds  ol  probable  Xcuophoni  species  are  also 
found  in  a  fault-bound  outcrop  of  ]iebbly  coarse-grained 
sandstone  o\erlookiu'j;  the  lower  iii'o  lea  \alle\  i  Figure 
1  1.  The  molluscan  fauna  associated  with  the  molds  sug- 
gest a  late  F.ocene  age  and  hence  assignment  to  the  Par- 
acas ( )ioiip, 

NwiDM)   {■"i  IKMVTION.  (  KNTRM,  (jllI.K 

Tile  Navidad  Formation  (Darwin.  1846)  is  known  for  a 
rich  molluscan  fauna  that  was  last  re\iev\'ed  extensively 
by  Philippi  (18S7'.  Sediment  deposition  as  indicated  b\- 
microlossils  ranges  from  intertidal  to  outer  shell.  Most 
of  the  spi'cinu'iis  of  Xcnopliorn  descrilied  iiere  have 
been  recovered  from  gra\  deepwater  siltstone  tliat  today 
forms  the  intertidal  platform  at  Pnnta  Perro  (Fignre  2). 
These  deposits  have  been  datetl  with  foraininii'ers  hv 
Dremel  (in  Herni.  19(i9.  p.  71  )  as  Power  Miocene  (Bnr- 
ch<falian>.    However.    Tsnchi    et    al.    (1990)    and    ll)araki 


Page  72 


THE  NAUTILUS,  \'ol.  116.  No.  3 


75°35"W 


~^ 


.1  ^.d^i  i 


75  "W 


PANAMERICAN 
HIGHWAY 


Figure  1.     Fossil  loculitics  lor  Xcnopluini  ctiiditiiH-rii  in  the  Pisco  Basin  ot  Pc 


(1992)   lia\v   poiiitcil   out    thai    an    Uppci'   Midfciic   acjc 
(Tortoiiiaii)  is  nKirc  likcK. 

.■\cklitioiial  spciiinciis  lia\c  hccn  colhilcil  ihhIIi  iiI  llic 
Ri'o  Hapel  and  Iroiii  Malanzas.  l)()tli  also  in  the  aioa  ol 
Na\idacl  (Fissure  2).  Tlic  scdimciils  ol  liicsc  localitirs  arc 
believed  to  he  (•(intrrniiorancoiis  with  tliosr  ol  I'nnia 
Perm. 

.\Iii,i.()\f;ri';  Foumaiion.  Pkmn.sula  Ahai'co.  sen  iii- 
cjENiHAi.  (.'im.i-: 

Tlie  .Vlillonirne  P'onnation  has  heen  defined  from  coi'es 
drilled  on  Peninsula  .\rauco.  It  consists  ort^nix-  sillstones 
dated  a.s  Kocene  (e.g..  Garcia.  1968).  .\\\  similar  sill- 
stones  of  this  area  usnallv  ha\e  heen  eonsidcicd  to  he 
long  in  this  f(jrniation.  Ilovvex'er.  analvsis  ol  tlu  niollns 
can  (anna  suggests  a  .Miocene  age  at  least  loi  pail  ol 
tlie.se  .sediments  (.S.  Nielsen,  unpuhlished  dataL 

The  Strlldria  specimen  descrilietl  herein  was  loniid  in 
a  concretionar\-  nodule  from  a  coastal  cliff  with  e\po 
sures  of  gray  siltstones  to  the  east  of  Pnnta  Millongne 
(Fignn^  2).  These  concretionan-  nodules  are  washed  Iree 


h\  the  tide  and  iisnalK  \ield  specimens  of  the  crah  ('iiit- 
ccr  finiiii(mii\  I'hilippi,  ISS7.  Because  this  localitx  has 
not  heen  d.iled  until  now.  the  age  of  the  Stelldria  rc- 
ni.iins  nneert.iiu. 


MATFHi Ai.s  wi)  Miriiions 


.Specimens  deselihed  in  mentioned  in  this  stud\  .ne  de- 
liosiled  in  the  (dileetlons  ol  die  lollowinij;  miiseiims;  l)e- 
partameiilo  de  I'.ileontolo'^ia  de  \  eitehiailos.  Miiseo  dc 
liisloria  Natural  de  la  Uni\crsidad  de  San  Marcos.  Lima. 
Peru  (MIISM  IN\'):  Departamento  de  Paleoutologi'a  de 
jnxcrtehrados.  Miiseo  Nacional  de  llistoiia  Natural. 
Santiago  de  ( liile  (SCO. PI):  and  Senckeuhert;  .Museum. 
Fr.uiklnrt,  (a'rniau\-  {SMP").  Photographs  were  taken  iis- 
iiil:,  a  Leicalle\  SLil  camera.  Images  were  scanned  Irom 
lllord  LP  I  \2~>  hlack  and  while  .'li  mm  negatives  using 
.III  \eei  SeanW'if  272()S  film  seaimer  and  processed  with 
Photoshop  (i.O. 


S.  N.  Nielsen  and  T.  T.  DeN'ries.  2002 


Fatie  73 


Figure  2.     Fossil  localities  tor  Xriiniiliiii'ii 
Chile. 


piiuliiKir  (\a\i(la(l  arcal  and  StvUiiria  kririurhdrtholili  (Peninsula  Aranco'  in  Central 


SYSTEMATIC  PALE0NT0L0C;Y 

FaniiK  Xenoplioridae  Philippi,  LS53 

Cienus  .\V/(()/)/)(i;v/  Fischer  xon  Waklheiiii,   ISO? 

Type  Species:  Xcnopltoni  l(ic\:i'^(itii  Fiselier  \iin  W'ald- 
lieiin,  1S07  (  =  Twchns  coHcliijluiphonis  Hoi-n,  ITSO). 

Xeru)phom  cardHi'UTii  new  species 
(Figures  S-U) 

Diajjnosis:  Moderate  size,  moderateK  tall  spiic;  base 
witliiiut  spiral  sculpture:  dorsal  surtace  with  weakly  op- 
isthocliuc  tci  spiral  irri"j;ul,u'  lirai'. 

Description:  Diameter  up  to  ofi  luiu.  Spire  au^le  70- 
yo"";  whoils  and  spire  Hat-sided  to  \er\  slightlv  coii\e\. 
Protocouch  poorK  preser\'ed.  Base  Hat  lo  sli'^hllx  con- 
cave; neither  spiral  nor  suhspiral  sculplure  i  lines, 
threads,  rugae)  evident;  colahral  giowtli  lines  ;iu(l  ridges 
present.  No  unihilii'us  in  :idiills:  none  ixidcnl  in  ju\('- 
niles;  thick  columella.  Dorsal  suilace  wiHi  wvaV  prosoe- 
line  growth  lines  and  coarse,  w;i\\  lir;ic  ihal  arc  weakly 
opisthocline  to  irregularK'  spiral.  No  prosocliue  nor  op- 
istliocline  axial  cords.  Cemented  objects  e\enly  spaced. 


about  se\en  pei'  w lioil.  increasing  in  size  on  later  whorls; 
less  than  oO*^-?  ol  shell  co\'ercd.  .Ajiertnre  unknown. 

Type  Material:  iiololxpe  S\ll'  :523()39  (figures  3-5), 
height  17  nun,  dianielcr  .»(i  niui;  o  parat\pes;  SMF 
.323040.  heighl  Id  uini.  diameter  :5(l  mm.  S.\1F  323041 
(figures  fi-S).  height  l.3.."i  mm.  diametei-  22. .^  nun, 
MUS.M  1N\'  1  ifignres  Oil),  height  21  mm.  diameter 
.32. .5  umi.  MUSM  INN'  2.  height  I.t  nun.  diameter  2fi.5 
nnn,  .ML'SM   IN\   .).  heiii;ht   |.>  nun.  diameter  21.5  nun. 

Tyjie  L<»ealit\:   D\   (S.5I-S.  northwest  of  l^)ma  (.'uesta 

Chilcatax.  about  1  km  north  of  Comotrana-C:arluias  road, 
about  5  km  east  Pla\a  (,'arhuas.  in  ridge-lorming  sand- 
stone bed,  120.5  m  in  measured  section.  14°l  I'Ofi"  S. 
7fi°0S'I7"  W  iPunta  Ciranik^  1:100.000  (luadrausilei. 

El-Miioloj^y :  Named  alter  ('anliln.  the  bi\al\c  most 
often  cemented  to  tliis  sjiecies.  and  gc;y;.  the  Latin  loot 
signif\in<4  'to  beai"  or  carrv. 

Occurrence:  Otuuia  l''ormatiou.  Oligoccne.  between 
Paracas  and  Bahi'a  de  la  Independencia.  I'cru.  I'ossibiy 
(rom  the  uppermost  Foc'cue. 

Discussi()n:  Specimens  ol  \tiHi}>hora  canliti^erfi  new 
species  dilter  in  several  respects  from  those  of  X.  roti- 


Paw  74 


THE  NAUTILUS.  \  ol.  1 16.  No.  3 


Figures  3-12.  Tfrtian  XiiKipluini.  3-11.  Xciiophora  canlitifit'iu  ni-w  .spciir.s.  ;}-.5.  I  l(il()l\])r.  .S.\ll'  .32:)(1.)U.  li(ii;ht  IT  iiiui. 
ciiameter  36  niin.  6-8.  Parahpe  SMF  323041.  lK•i^ht  13.5  mm,  diameter  22.5  mm.  9-11.  Parahpe  .Ml  SM  IW  1.  htiglit  21  mm, 
diameter  32.5  mm.  12.  Xenophora  pmtlmac  ne«-  speeies.  Parat\pe  S(;().l'l  5991.  (lianictcr  154  mm. 


S.  \    Nielsen  and  T.  J.  DeNries.  200:; 


Pa<fe  75 


clujliophora  (Bom.  1780).  a  species  with  a  fossil  record 
that  may  e.xtend  back  to  die  Eocene  and  wiiich  is  pres- 
ently found  off  the  coast  of  bodi  eastern  and  western 
North  and  Central  .\merica  (Ponder.  19S3!.  The  latter 
species  is  characterized  by  rugose  cords  on  tiie  dorsal 
surface;  wa\y  lirae  that  are  distincth'  opisthocline;  base 
with  weak  spiral  sculpture:  an  umbilicus  in  ju\  enile  spec- 
imens: and  a  thin  columella  and  umbilical  callous  in 
adults.  Specimens  of  .V.  carditi^cro,  in  contrast.  ha\e  nei- 
ther rugose  spiral  cords,  stronglv  opisthocline  lirae.  nor 
spiral  sculpture  on  the  base,  and  ha\e  a  thick  rather  than 
thin  columella. 

Xenophora  carditigera  most  closeK'  resembles  .\'.  ficinin- 
<l)  Beu.  1977.  an  earK-  to  middle  .Miocene  species  from 
New-  Zealand  diat  may  be  part  of  a  lineage  diat  includes 
X  prognata  (Finlay,  1926)  (late  Eocene  to  middle  Mio- 
cene) and  .\'.  ncozehinico  Suter.  1908  ie;ul\  Pliocene  to 
Recent).  .Specimens  oi' X.  fleniiiigi  are  iilso  lUoderateK' tall 
and  straight-sided,  lack  an  umbilicus.  ha\"e  weakl\-  devel- 
oped to  obsolete  spiral  sculpture  on  the  base,  and  are  onl\- 
partialK'  covered  b\-  cemented  debris  diat  increases  in  size 
towards  the  apeitm-e.  The  princip;il  difference  between  X. 
carditigera  luid  .V.  flcmingi  is  diat  dorsal  lirae  on  the  latter 
are  distincd\'  opistliocline. 

Xenophora  pnulinac  new  species 
(Figures  12-16) 

Diagnosis:  Shell  large.  umbiBcus  closed,  whorl  outline 
concave.  Subsutural  ramp  reaching  halR\a\  onto  previ- 
ous whorl.  Dorsal  surface  between  cemented  objects 
with  coarse  wa\T  hrae  and  prosocHne  axial  growth  lines. 

Description:  Shell  large  (liolot\pe  diameter  132  mm. 
height  62  mm '.  with  depressed  spire  (angle  about  105°), 
peripheral  flange  unknown.  Protoconch  unknown.  No 
umbilicus  in  adults  (no  juveniles  known),  whorl  outline 
concave  due  to  subsutural  ramp  reaching  about  halfwav 
up  onto  previous  whorl.  Dorsal  surface  between  ce- 
mented objects  with  coarse  wav-\-  lirae  as  well  as  prosoc- 
line  axial  growth  lines.  Base  weaklv  concave,  sculptured 
with  low,  narrow,  irregular,  collabrid  growth  lirae.  .At- 
tached camouflaging  objects  seem  to  include  either 
high-spired  gastropods  or  concave-side-up  bivalves  lup 
to  50  mm  wide).  Basal  apertural  lip  regularlv  and  shal- 
lowiv  cursed,  .\perture  unknown. 

T\pe  .Material:  Holotvpe  SMF  323042  (figures  13- 
14),  height  62  mm,  diameter  132  mm.  Punta  Perro; 
paratopes  SCO. PI  5991  (figure  12),  diameter  154  mm. 
Punta  Perro.  SGO.Pl  5992  (figures  1.5-16),  spire  frag- 
ment, height  16  nun.  Matanzas. 

Type  Locality:  Intertidal  platform  at  Punta  Perro, 
central  Chile. 

EtyTnolog\:  Named  after  Paulina  S.  N'asquez  IlliUU-s. 
friend  and  colleague,  who  found  part  of  the  tvpe  materi;il. 

Occurrence:  Navidad  Formation.  Tortonian.  Navidad 
area,  centred  Chile. 


Discussion:  Xenophora  patdinae  new  .species  differs 
from  most  other  species  by  its  large  size.  The  onlv  sim- 
ilarly large  species  is  the  Eocene  to  Miocene  New  Zea- 
land Xenophora  prognata  ( Finlav.  1926)  isee  Beu.  1977) 
from  which  it  differs  in  having  concave  rather  than  con- 
vex whorls,  formed  by  a  sidjsutural  ramp  reaching  about 
halfxvav  up  onto  previous  whorl. 

Previouslv.  Tavera  il979)  stated  that  his  Trochita  gi- 
gantea.  also  from  the  Navidad  Area,  might  prove  to  be 
a  Xenophora.  However,  Tavera  (1979)  provided  no  di- 
agnosis to  separate  his  species  from  other  taxa,  and  he 
did  not  figure  the  specimen.  Conse<juentlv  the  name  is 
considered  imavailable  under  ICZN  Article  13.1.1. 

Genus  Stelkiria  Schmidt.  1832 

Tvpe  species:  Trochus  so/c/n.s  i>iime.  1764. 

Stellaria  hriegcrl)artholdi  new  species 
(Figures  17-20) 

Diagnosis:  Spire  short,  with  narrow  peripheral  flange 
divided  into  prominent,  blunt  digitations.  No  umbilicus. 
Base  lightlv  convex,  with  distinct  collabral  growth  lines. 

Description:  Short  spire,  periphen  divided  into  about 
10  prominent  blunt  digitations.  No  umbilicus.  Whorl 
outhne  and  sculpture  unknown,  because  original  shell  is 
dissolved  except  for  peripherv  and  digitations,  but  out- 
line appears  to  be  weaklv  convex.  Base  shshtlv  convex, 
with  distinct  collabral  growth  lines. 

Holotype:  SGO.Pl  5993  (figures  17-20).  height  24 
mm,  diameter  with  attachments  60  mm.  spire  angle  9.5°. 

T\pe  Locality':  Northeast  of  Plava  Millongue.  Penin- 
sula .franco,  south-central  Chile. 

Etvmology:  Named  after  Rolf  Kriegerbarthold.  who 
did  the  difficult  preparation  of  this  specimen. 

Occurrence:  Northeast  of  Punta  Millongue.  Penin- 
sula .\rauco.  south-central  Chile. 

Discussion:  Stellaria  kriegerbartholdi  differs  from  all 
other  species  of  this  genus  in  having  a  clo.sed  umbilicus. 
The  tvpe  species,  S.  solans,  has  tubular  spines,  a  feature 
that  cannot  be  observed  in  S.  kriegerbartholdi.  S.  krie- 
gerhariholdi  most  resembles  the  Oligocene  to  Pliocene 
S.  testigera  (Bronn.  1831).  with  its  two  Recent  subspe- 
cies, in  having  a  stronglv  digitate  peripheral  rim.  -S.  tes- 
tigera lived  in  the  Mediterranean  and  spread  to  Atlantic 
.Africa  and  the  Gulf  of  .Aden.  .As  suggested  by  Ponder 
(1983).  -X  testigera  could  have  evolved  from  the  Eocene 
S.  conica  (Dall.  1892'  from  Mississippi,  which  also  might 
be  regarded  as  ancestral  to  .S.  kriegcHjariholdi.  S.  conica 
has  the  umbilicus  edinost  obscured  bv  the  parietal  callus 
.Mc.Neil  and  Dockerv.  1984).  The  completely  closed 
umbilicus  would  also  justilv  inclusion  of  this  species  in 
Xenophora.  suggc>sting  that  this  species  or  species  line 
lost  its  camouflaging  habit  and  evolved  from  a  difTerent 
ancestor  than  Stellaria  s.s.  However,  this  view  is  not  fol- 
lowed here. 


Pasie  76 


THE  NAUTILUS.  \nl.  llfi.  No.  3 


Figures    I3-l(i.   Xnwphora  pauluwr  ur.v  spvirs.    i:?-14.   llolulxp...  SMI'  o23()l2.  lu-,^1,1  h2  nnn.  clianu.l.-i    1:52  un 
Paratvpc-  S'- .>  M'l  3i)92.  spire  IVauiiu'iit,  lici^^lil  Hi  iiiin. 


1 5- Hi. 


Remarks:  '\\,<  di.isiii'  .I  Ibi-  Stclltiriti  'j;i\fii  In  I'diuKt 
(19S3)  lias  to  III-  cnicndcd  to  indnclc  spct-ics  witli  a 
closed  iiinhiliciis  like  S.  kiiciicrhdrllioldi.  Ponder  (19S3) 
noted  that  die  placeiiieiil  ot  S'.  icsti<iii'ra  "is  iml  eiini- 
pieteK  satislacton".  a  eoiniiiciit  wliicli  is  excii  nunc  ap- 


prnpnalr  Inr  S,  kiii-^nhinlliohli  ll  does  nut  sc.-m  jus- 
tilird  to  creel  a  new  '^einis  lor  tins  spet'ies  heeanse  a 
closed  nnihiliens  is  also  preseni  ni  \, iinpIiKril  and.  tlieie- 
lore.  seems  lo  he  an  old  in\entinn  ol  tlie  lamiK.  Cliar- 
aelers  separatni'j,  ,S7,  //r;/;f/  Irom  Xiiioplii'm  are  reduced 


S.   \.   Xiclsc 


T.  1    i:)f\ries.  2002 


Wm- 


Finiirt's  17-20. 

(iO  iiini. 


Slillariti  kiif^crhiiiilioltli  new  species,  llolofxpe,  SCiO.I'l  .5SJ!).).  lieij^hl  24  iinji.  diaiiieter  witli  .ittaeliiiieiits 


to  tile  preseiiee  (il  spiiu-s  or  (limitations,  an  expandi'tl 
peripheral  flaiiij;e  and  a  iieaiK  sniootli  dorsal  surlaee. 

CONCLUSIONS 

The  Cenozoic  Nenophoridae  ul  Cilnle  and  I'eiii  elose  a 
considerable  biosreographic  gap  in  the  histon  ol  the 
taniiK'.  Oligoceiie  records  of  XcitDjtliora  are  lew  but  the 
genus  was  alread\-  established  in  Australia  (Ponder  19S.'3) 
and  New  Zealand  (Ben  1977)  in  the  Eocene.  In  the  Mio- 
cene the  genus  was  widespread  in  the  Indo-Paeific  {Re- 
gion with  tliiee  species  also  picsenl  ni  the  CaiiMiean 
Sea.  Toda\^  onK  the  t\pe  species,  X.  coiulnilii'phnni.  re- 
mains in  the  Aniericas  (Ponder  198.3). 

Our  t^\(l  species  ol  XmopJiDni  seem  closeK  allied  w  ilh 
New  Zealand  species.  Similarities  between  launas  Ironi 
New  Zealand  and  Argentina  ha\e  recentK  been  indicat- 
ed b\-  Ben  et  al.  (1997).  Our  records  ol  Xciiojiliiira  show 
that  there  are  even  more  connections  witli  Pacific  South 


.Vnieiiea.  a  laet  whieh  is.  ol  course,  not  surprisin'4.  Moil- 
tl'.ius-l-'acilic'  alliuities  can  be  expeclcd  Irom  lurlher 
studies  ol  Pacific  South  .Amerii'an  launas. 

Stclliiriti  ctiiiicd  Irom  isocene  beds  ol  .Vlississijipi  ma\' 
be  an  ancestor  of  .S,  kric^crlxirlliold't  but  has  an  open 
umbilicus  like  modi'in  sjiecies.  i'onder  (198.3)  discussed 
,S.  Icsllis.<'i''i  as  a  possible  carK  ollshoot  while  Stcllaria 
was  still  close  to  \ciioj)liorii.  which  ma\  also  be  true  lor 
.S.  krii'<H'rh(irthol(li.  llowc\cr.  characters  ol  (lie  known 
Stcllaria  species  suggest  that  the  lossil  record  ol  (his  ge- 
nus is  \(:'r\  IragmenlaiA. 

ACKNow  i.i:i)(:mi;x  IS 

Klaus  Handel  is  diaiiked  for  companx  in  the  fii'ld  and 
discussion  on  (Jiilean  geoloin  and  bioloi^ol  Xciiophora. 
|ose  II.  Peal.  .\lau  Ben.  and  an  anonxnions  reviewer 
made  \aluablc  counnents  which  heliied  to  improve  the 
manuscript.  Kieklwork  ol  SNN  was  linaucialK  supported 


Pa«'  7S 


THE  NAUTILUS.  No 


Ifi.  No.  3 


bv  the  Deutsche  Forsdiuiigsgeiiieinsehaft  (DFG),  grant 
Ba  675/25.  Fieklv\ork  ofTJDV  wa.s  financed  In-  liimseli 
and  in  part  bv  a  Fulbright  Senior  Scholarsliip. 

LITERATURE  CITED 

Biindel.  K.  199.3.  Caenoga.stropoda  during  .Mesozuif  tiniL'S. 
Scripta  Geologica,  Special  Issue  2:  "-.56. 

Beu.  A.  G.  1977.  New  Zealand  Cenozoic  gastropods  of  the 
genus  Xenophora  Fischer,  1807.  Journal  of  the  Ro\al  So- 
cietv  of  New  Zealand  7:  229-24 1 . 

Beu,  A.  G.,  M.  Griffin  and  R  A.  Maxwell.  1997.  Opening  of 
Drake  Piissage  gatewa\  and  L^ate  Miocene  to  Pleistocene 
cooling  reflected  in  Southern  (Jccan  nioilnsean  dispersal: 
evidence  from  New  Zealand  and  .Xri^entina.  Tectonoph\s- 
ics  281:  83-97. 

Darwin,  C.  1846.  Geological  (iliscnations  on  South  ,*\merica. 
Smith,  Elder  &  Co.,  London,  279  pp 

DeX'ries,  T.  J.  1998.  Oligocene  deposition  :uiil  ('enozoic  se- 
quence boundaries  in  the  Pisco  Basin  (Pern).  Journ:il  ol 
South  .American  Earth  Sciences  11:  217-231. 

Dunbar  R.  B.,  R.  C.  Mart^-,  R  A.  Baker  1990.  Cenozoic  marine 
sedimentation  in  the  Sechura  and  Pisco  basins,  Peru.  Pa- 
laeogeographv,  Palaeoclimatolog)',  Palaeoecologv  77:  235- 
26L 

Garcia  A.,  F.  1968.  Estratigrafia  del  Terciario  de  Chile  C:entral. 
In:  Cecioni,  G.  (ed.):  Svmposio  sobre  el  Terciario  de 
Chile,  Zona  Central.  Andres  Bello,  Santiago,  pp.  25-57. 


Herni,  D.  1969.  Marines  Pliozan  und  Pleistoziin  in  Nord-  und 
Mittel-Chile  unter  besonderer  Beriicksichtigung  der  En- 
h\icklung  der  Mollusken-Faunen.  ZitteUana  2,  159  pp. 

Ibaraki,  M.  1992.  Planktonic  Foraminifera  from  the  Naxidad 
Formation.  Chile:  their  geologic  age  and  paleoceano- 
graphic  implications:  pp.  91-95.  In:  Ishizaki.  K.  and  T. 
Saito  (eds.):  Centenan  of  Japanese  Micropaleontolog\-. 
Terra  Scientific  Publishini^  Company,  Tok\o. 

Kreipl,  K.  and  A.  Alf  1999.  Recent  Xenophoridae.  Conch- 
Books.  Hackenheini.  148  pp. 

McNeil,  F.  S,  and  D.  T  Dockerv.  III.  1984.  Lower  Oligocene 
Castnipoda.  Scaphopoda,  and  Cephalopoda  of  the  Vicks- 
burg  (iroup  in  Mississippi.  .Mississippi  Department  of 
Natural  Resources.  Bureau  of  Geolog\-  Bulletin  124.  415 
pp. 

Philippi,  R.  A.  1S87.  Los  losiles  terciarios  i  cuartarios  di-  (^liile. 
FA.  Brockhaus,  Leipzig.  256  pp. 

Ponder  W.  F.  1983.  Xenophoridae  of  the  world.  The  .Australian 
.Museum  Memoir  17:  126  pp. 

Ta\era,  J.  1979.  Estratigrafia  \'  paleontologi'a  de  la  F'ormacion 
Navidad.  Provincia  de  Colchagua.  Clhilc  iLat.  .30°50-34° 
S).  Roletin  del  Museo  de  llistoria  Natural  (diile  36: 
1-176. 

Tsuchi,  R.,  T  Shuto,  T  Takavama,  I.  Koizmni,  A.  Fujiyoshi,  .M. 
Ibaraki  and  Martinez-R.  Pardo.  1990.  Fundamental  data 
on  Cenozoic  biostratigraphv  of  Chile.  Supplement.  3. 
Punta  Perro  section  north  of  Navidad.  central  Chile.  Re- 
ports of  Andean  Studies,  Special  N'olume  3:  70-71. 


THE  NAUTILUS  116(3):79-S.S,  2002 


Page  79 


Redescription  of  the  genus  and  species  Hctcrova^iiiiiui 
Jimaijana  (Lesson,  1830)  (Gastropoda:  Soleolifera: 
Veronicellidae! 


Jose  \>'illilialclo  Thome 
Siizelc  Rodrigues  Gomes 
Kosane  Soiiza  da  Silva 

Lalidratinii)  de  Malacologia, 
Faculiliuk-  de  Biociencias.  PUCRS 
9()69()-9()()  Porto  \h'2\v 
BRAZIL 
tlioniej\\(a'pii(.'is.hr 


ABSTRACT 

Based  on  the  examination  of  58  speeiinens  deposited  in  xarioiis 
institutions,  we  redeseribe  the  genus  ll<tcroi:a'^inina  Kraus, 
1953,  considering  it  inonot\pic  and  with  its  range  restricted  to 
Penj.  The  t\pe  species,  H  linuiiicniii.  ( l>esson.  1S30)  is  also 
redescrilied,  and  a  neoUpe  de.signated  for  the  species.  The 
anatonn-  ol  tlie  herniaphioditic  region  near  tlie  female  genital 
pore  is  emphasized  as  characteristic  ot  the  genus:  the  copula- 
tion bursa  is  elongatetl'spheroid.  tapering  into  a  distal  elonga- 
tion tliat  tolds  and  joins  tlie  bursa  duct  toward  its  base,  receiv- 
ing the  junctor  duct  at  the  tip  of  the  bursa.  The  rectum  pen- 
etrates the  integument  near  the  female  genital  pore  i'liiic  is 
no  penial  glaud  and  no  accessor\'  gland.  The  aualum\  ol  the 
anterior  male  gemtal  s\stem  cliaracterizes  the  species:  smooth 
penis,  with  short  cvlindrical  \erge  that  intumesces  slighth  in 
its  medial  region,  forming  expansions  on  the  sidis  tli:il  mark 
the  beginning  ol  the  glans;  tlie  penis  is  lougei  llicn  Hie  xerge, 
and  tapers  in  its  distal  portion.  There  is  no  spatlia.  All  pertinent 
hibliugiapliN  is  discussed  :ind  the  SMion\iui/.ation  of  \'«g/;/i/i(/ 
(hIIuiiii  I  liilliiKinn.  UJ2T.  is  justified. 


IXTHODUCTION 

The  WronicelHdae  are  terrestrial  sings  with  worldwide 
ilistrihiition,  stiii  poorK'  knowii,  and  ot  uiKcrtain  position 
witliiii  tlie  Ca.stropoda  (Hoffmann.  1925:  ii\nian.  UKiT: 
Thome,  19SSa,  1993;  Tillier  et  al„  199fii.  Among  the 
genera  iiielndcd  in  the  famiK'  (Hotfmami,  1925:  i'orcart, 
1953;  'flioiii('\  1975),  t^\o  sliare  the  lack  ol  :i  penial 
gland,  a  stiiietme  foinied  h\  a  papilhi  ;ui(l  a  varied  iinni- 
her  ol  tubules,  wliu-ji  lies  alongside  the  penis  and  lias 
swstt'inatii'  importance,  altliongli  its  tmietion  is  vet  not 
understooil. 

Ol  the  hvo  genera  ol  the  \  eionieellidae  lacking  the 
penial  gland,  one  occurs  in  ,\friea.  \'(i<^iniiHi  Simrolh. 
1S97,  and  the  otliei',  Hctcwva^imtid.  is  I'estrieled  to 
Peru.  Sinn-otli  characterized  Vfi'jjiiind  simpK  1)\  the  ab- 
sence of  the  penial  gland,  a,s  opposed  to  the  genus  Va- 


"iiuild  l^ertliold,  bS2' 


:i  sv  noiiv  111  () 


\  aiiiiiithts  Fernssac, 


1S21).  vvliicli  exliibils  (bat  siriietiire.  In  |iroposing  the 
genus,  Simidtli  !lS!)7'  based  it  on  the  e\ainiiialion  of 
two  specimens  without  speeilie  names.  Oiilv  later,  thai 
author  iSimrofh.  I9l.it,  examining  the  same  two  speci- 
mens, di'scribed  iheiii  as  V  counulli  and  \!  loffu'nsis, 
both  originating  Iroiii  the  same  West  .Mrican  loealitv.  I?iit 
tliev  differ  In  the  presence  of  glandular  jirotiibeiances 
on  one  side  ol  the  |)eiiis  sheath  in  \  coiiniilti.  ami  bv  a 
smooth  sheath  in  \'  /()g<ie)is7s 

In  his  disciissioii  of  the  iiomenciatiire  of  the  \  eroiii- 
cellidae.  Baker  (1925)  designatecl  \'aiiiiiiiia  rcnnuUi  as 
the  t\pe  species  of  Vd'^inhui.  liollmami  (  1925)  accepted 
\'  roiinnlti  and  sv  nonv  mi/ed  W  lofiocusis.  maintaining 
that  the  character  used  bv  Siinroth  (1913)  would  be  in- 
sufficient to  diflerentiate  them,  in  that  he  bad  onlv  fvvo 
spi'cimens,  both  collected  in  the  same  localitx.  Ilotf- 
maiin  (1925)  included  the  genus  in  his  then  new  siih- 
tamlK  .Meisenheimeiiinae,  with  the  following  diagnosis: 
"dark  notum,  vvcaklv  pointed;  genital  jiore  almost  at  inid- 
lengtli  ol  the  bvponolum:  with  (he  anterior  lobe  of  the 
digestive  gland  behind  the  anterior  loop  of  the  intestine; 
with  the  pedal  iiei-vcs  together  for  a  short  stretch,  par- 
allel and  then  diverging:  with  a  needle-shaped  penis  and 
;i  basal  ringed  tbiekeiiing:  without  penial  gland". 

I  lolliiKiiiu  d927)  desi-iibed  Vaiiiiiiiiti  odiuwri  based 
on  a  single  specimen  of  unknown  origin,  but  stating  that 
it  was  certainlv  from  .Mriea,  despite  the  lact  that  W/g/- 
nina.  known  oiiK  from  .\liica,  vv:is  onlv  recorded  once. 
Without  discussing  the  smioiimiis  he  had  proposed  in 
1925,  he  differentiated  \'.  (Hlliiicri  from  V.  ioffjcnsis  by 
the  shape  of  tlie  penis  (also  different  from  the  gemis- 
levcl  inoiphologx-  lie  had  proposed)  and  respective 
sheath  ami  asi^'cts  of  the  copulation  bursa  and  con- 
necting ducts,  lie  adds  tli;it  in  his  species  the  lobe  of 
the  digestixe  gland  is  in  front  ol  the  iinterior  loop  of  the 
intestine  (also  dilleri'iit  Iroiii  the  jiroposed  gemis-levcl 
inoiphologx  1. 


Page  80 


THE  NAUTILUS.  \'ol.  llfi.  \( 


[•"ore-art  (1953).  in  his  monograpli  on  African  Wroni- 
ccllidae,  consiclcred  Vii<iitiiua  as  a  suhs^cnns  of  Pscudotc- 
roniccUa.  stating  tiiat  it  dilfcrs  from  tfic  otficr  snfigcncra 
inclnck'd  in  tlu-  gcnns  1)\'  tfu'  characteristics  of  tfic  cop- 
nfaton'  organs  and  tfic  prcscnci-  of  canafs  and  ducts  iic- 
tween  tlu-  rectniii  and  the  int'tcr.  witiiont  discussing,  at 
neither  tlie  gen\is  nor  llie  sntigenus  fevel,  the  absence 
or  presence  of  tlic  pciiial  glantf.  Forcart  (1953)  agreed 
with  the  s\'non\ni\  of  Hoffmann  (1925)  and  considered 
Rscii(li)rcr()iiicclla  lA'd^iiiiiui)  roiiriidti  as  a  \alid  name. 
W  itfiont  (fiscussion  an(f  witfiont  examining  specimens, 
he  afso  atfdecf  PsciKhiirroiiicflla  [\'ap,iiuiia}  odhncii 
(Hoffmann)  to  fiis  list  as  from  "prol)alil\'  Africa".  He  also 
proposed  a  new  species,  PseudovcmuiccHa  (Vaf^inina} 
diipoiii.  obsening  that  this  would  differ  from  the  others 
in  the  subgenus  l)\  the  presence  ot  the  penial  gland 
(which  in  fact  sets  the  species  apart). 

Kraus  (1953a)  proposed  the  ta.\on  IhicroiniiiniiKi  as 
a  subgenus  of  Vaginina,  baseil  on  the  description  of  \'(i- 
ginina  (Heterovaginina)  penivicnia,  from  Lomas  tie  .\to- 
congo,  near  Lima,  Peru,  and  included  \'  odhiich  due  to 
the  similarities  of  the  two  species.  He  excluded  V.  foii- 
radti  !)ecause  the  anterior  intestinal  loop  is  anterit)r  to 
tlie  digestive  gland,  the  copulation  bursa  lias  a  long  duct, 
and  the  penis  has  a  much  different  shape.  Kraus 
(1953b),  without  anv  reference  to  Baker  (1925)  or  to 
Hoffmann  (1925),  but  mentioning  verbal  contact  \\itli 
Forcart  (Basel,  Switzerland)  and  based  on  the  work  of 
Degner  (1934)  on  tlic  analoni\  ol  W  lugociisis.  raises 
Heterovaginina  to  the  genus  le\el.  Kraus  stated  tliat,  af- 
ter examining  histological  sections  of  a  paratvpe  of  H. 
pcnnidiuL  he  found  it  had  a  cloaca  and  lacked  the  links 
between  the  rectum  and  the  tbnrtli  nreti'r,  thereb\'  dif- 
fering from  the  African  genus  that,  according  to  Degner 
(1934),  had  separate  openings  for  the  rectum  and  the 
fourth  ureter  that  were  connected  to  each  other  b\  a 
urethral  canal  and  5  urethral  ducts  (which  were  used  in 
the  description  \)\  Forcart  (195.3)).  Later  on,  Kraus 
(1954)  s\iion\nii/,ed  Heterovaginina  jieruviaiui  Kraus. 
1953,  with  Vagiindu\  liniai/uinis  Lesson.  IS.'IO.  consid- 
ering that,  according  to  peisonal  inloinialion  from  \V. 
Wevrauch,  "only  one  species  occuis  "  ni  the  t\pe  loealitx. 

Forcart  (1957),  after  examining  I'eruxian  specimens  in 
the  I'ield  Museum  of  Natural  lliston  (now  re-examined 
by  us),  found  them  to  represeni  \a'j,iiiiii)i  odliiicri.  Iml 
concluded  that  this  species  should  lie  included  m  the 
genus  Heter()va<iinina  because  il  oecurs  m  southern 
Peru.  He  thus  agreed  with  Kraus  (1953a).  allliou^li  lie 
did  not  slati-  so  directK. 

Thome  (1969)  redescribed  the  hololxpc  <)\  \agininii 
(Heter<n:aginiuti '  peniviana  Ki'ans,  1953,  depositc'd  in 
the  Seiickenberg  .Museum.  l'"rankfurt-am-Main,  williout 
discussing  the  .s\iion\inv.  The  copulation  buisa  is.  ae 
cording  to  Thome  :  1969),  sac-like  and  sessile  in  \:  i  //  ' 
peniviana  (which  now  we  know  to  be  present  oiiK  in 
juvenile  specimens  L  Thome  (1975),  in  a  revision  of  tlie 
American  g(>nera  of  \erouieellidae,  recogni/ed  and  re- 
describetl  (still  only  on  the  basis  of  the  juvenile  t\|ie 
specimen)  Heterovaginina.  keeping  il  iiionol\]iic  i //  j)e- 


nn  ianai  and  with  otcurrence  restricted  to  Pern,  lie  did 
not  mention  \    odiineri. 

Ill  addition.  Thome  (  19S4)  ri'described  the  liolot\pe 
of  //,  odiineri.  di'positetl  in  the  Natiniiistoriska  Kikmu- 
seet  ol  Stockholm.  Swetlen.  .According  to  Thome  ( 1984), 
the  cdpulatiou  bursa  is  s|iheroid.  seated  ox'er  a  thick  cv- 
lindrieal  uniform  duct  and  has  a  short,  slender  junctor 
diK't  that  penetrates  the  copulation  bursa  duct  at  niid- 
leiigtli.  An  inspection  of  its  illustration  (Figure  18,  p.  44) 
and  kiiowiedm'  that  the  material  was  damaged,  lead  us 
to  infer  that  the  (.(ipnlation  bursa  had  actually  been 
pushed  (.lown  along  its  duct.  Thus,  the  junctor  duct  is  in 
part  adlu'iing  to  the  bursa  duct  and  does  not  penetrate 
it.  'I"lu'  junctor  duct  must,  certainly,  penetrate  the  tip  of 
the  damaged  copulation  bursa.  The  penis  is  short  and 
broad,  with  a  short,  level/convex  verge  that  continues 
toward  tlie  spheroid  glans.  This  latter  is  trans\ersall\' 
widened,  witli  a  lateral  conical  protuberance  projecting 
from  the  le\el  face  of  the  verge.  Thome  et  al.  (1999) 
reported  a  preliminaiy  illustrated  redescription  of  the 
genus,  which  is  now  presented  herein  in  an  expanded 
format. 

During  the  examination  of  the  specmu'us  aiut  ol  tlie 
literature  to  define  the  species  of  the  Wronicellidae  oc- 
curring in  Peru,  several  specimens.  Iiotli  juvenile  and 
atliilt,  were  found  in  the  examined  collections.  These 
were  positixelv  identified  as  Heterovaginina  limaijana 
(Lesson.  1830).  Based  on  this  material  we  propose  the 
validitv  ol  the  genus  and  the  species,  with  their  respec- 
tive redescri|itioiis  and  the  designation  of  a  neotvpe. 

MATERIALS  AND  METHODS 

Fiftx'-eight  specimens  from  26  lots  were  examined,  from 
the  collections  of  the  Museo  de  Historia  Natural.  Univ- 
ersidad  Nacional  Mayor  de  San  Marcos  (MUS.M),  Lima, 
Peru;  Senckenberg  Museum  (SMF),  Frankfurt-am- 
.\laiii,  Germany;  Field  Museum  ol  Natural  Histon- 
(F.MNH).  Chicago.  USA;  British  Musenni  of  Natural 
lliston  iBMNH),  London,  England;  .\luseu  de  (aeucias 
e  Tecnologia,  Pontiticia  Unixersidade  C^atolica  do  Rio 
(;;rande  do  Sul  (MCP-PUCRS),  Porto  Alegre.  RS,  Brazil; 
-National  Museum  of  Natural  Histoiy  (USNM),  Wash- 
ington, DC,  USA  (Thome  et  al.,  1997)  and  Natnriiisto- 
liska  liikmnseet  (NRS),  Stockholm.  Sweden  (Thome. 
10841. 

For  studv  of  the  internal  anatomv.  the  specimens  were 
dissected,  starting  with  a  posterior  to  anterior  longitu- 
diiKil  ventral  incision,  follovvin-j,  llie  iiictliodologv  ol  'l1io- 
nie  and  I  ,i  ipcs  (19731  ;ind  siil)si'(|iient  iinprov  emeiits. 
Tlic  iiiaiidililc  (  =  j;ivv.  ol  sonic  .mlliorsl  and  radula  of 
two  specimens  lidm  lots  \1CP  6594  and  MUS.M  251.'Vb 
were  e\li;icled  inidci  the  slereomicroscope  and  later  ;in- 
;ilv/ed  ;ind  iiie;isiired  under  the  sc;iiiiimi,i  elcctiiiii  mi- 
croscope. The  ;iv('rages  ol  length,  width,  and  disl.inci' 
betxveen  the  region  of  greatest  width  :ind  the  anlerior 
lip  of  Ivvc'jitv  rac'hidiaii  teeth  ;ind  twentv  lateral  teeth 
vveic    c;ilciil;ited.     I'lic    anatomic,    generic    :iiid    spi-cifit'. 


].  w.  Tl  11)11  K-  ,i  III.  -loo-: 


Vim-  SI 


cliaiai-tt'ristifs  are  einpliasizrd,  Inllow  lipj,  I  loll  niaiiii 
(1925)  and  TlioiiK'  (1975.  19SSa.  I9SS1),  I99.)i. 

SVSTEMATICS 

Ck'iius  Hctcntiaoinina  Kraus.  1953 
Hctvrovd'^itibui  Kraus.  1953a:  fi.3-(S5. 

Type  Species:  VdiibiiiKi  1 1  hliini  ii^j^iniiui '  jirnn'uiud 
Krau.s.  1953a:  63-65  (=  Urlciiu  ii<j^iitiiiii  liiuiiiiuita  (Les- 
son. 1S3())). 

Diagncsis:  The  reetiiiii  peiietiates  (lie  iiite'j;iNiieiil 
near  the  female  t^enital  poi'e.  The  tliiii-tissiied  edpiilatioii 
bursa  is  spheroid,  taperiinj;  in  a  distal  leiiiitlieniuL;  that 
lieuds  and  joins  the  bursa  tluct  toward  its  base,  reeeivint; 
the  jiiuetor  duet  at  the  bursa  tip.  The  biirs:i  dnet.  nl 
iiioi'e  rit^id  tissue,  is  exlindrieal  and  e\treiiiel\  short  in 
the  juxeuile  aninial.  but  lengthens  in  the  adult,  bet-oin- 
in<r  an  eiontiated/spheruid  bursa.  \o  aeeessoi-x  inland  and 
no  penial  ^laml  are  to  be  found. 

Distrihiilion:      Kestrieted  to  I'eiu.  .South  .\iiieriea. 

Hiii-niniiiiitiiiii  liiiKii/diiii  (Lesson,  1830) 

\'irj^iiiiiliis  liiiiiiiiiiiiii\  LcsMiii.  1S30:  302-303.  471    pi    1  I,  fi<^s. 

I    I  \ 
Vifnimlii  liincdciiui  Deslia\es  in  Ferussae  and  Deshases.  1S19/ 

1S."j1:  V)(S'-9fr.  pi.  .SE.  fig.  11  (exact  copv  of  Lesson.  1S30). 
\'ii'j,iiiiiiii  ndhiirh   Hofhnann.    1927:   2(i.   fit^s.    II    II    'I'liciine. 

I9S4:  34-3fi.  figs.  14-19. 
Psi-itdovi  niidiilhi  (Va0inn(i'  (Hllnuir  l-'oitart.  1953:  (iO. 
Vii'^iiiiui    I  Ihtirdia^niiKii   jicnuKdui    Kraus.    1953a:    ^-(iS: 

Tlioine.  1969:  357.  figs.  29-30.  4S 
\'ai2iniiia  \HcterdV(i0iiiiui  <  oilhitcii:  Kraiis,  1953a:  fi3-fi5. 
Hetcrovii0iiiii/i  liiiKii/aiia:  Kraiis.  1954:  S2-S3. 
Hctcnivdi^iidiiii  dilhiirri:  Forcart.  1957:  9(S. 

Diagnosis:  Penis  smooth,  short  exliiidrieal  \erge  that 
intinnesees  a  little  bexoiul  the  middle  ol  its  length,  lonie 
ing  expansions  on  the  sidt's  that  mark  the  beginning  ol 
the  glaiis.  tdaiis  longer  than  th(^  \-erge.  tapering  in  its 
distal  portion;  generalK'  euncd.  There  is  no  spalha. 

Redeseription:  Extcnuil  Aiuiliddi/  A  large  pmiioii 
ol  the  I'xainineil  material  is  externalK  xen  diseoloicd. 
However,  the  color  pattern  on  tiie  iioliini  sliowed  ihe 
tollowing  general  characteristics:  specimens  with  light 
chestnut  colored  notuin  with  irregnlaiK  distributed 
black  dots  and  splotches.  ue\crtlieless  ciearK  showing 
two  longitudinal  black  lines  that  deliinit  a  lo/enge- 
shapeil  mid-region.  Between  these  two  lines  a  fine  light 
streak  occurs  centialK  in  the  uotiini.  starting  :i|iail  Ikhm 
the  anterior  pt'riiiotiiin  at  about  '4  ol  llie  wa\  down  the 
lull  length  of  the  animal  and  cont  inning  to  I  hi'  n'ar  witii- 
out  reaching  the  posterior  perinotiim.  (.Veragi's  ol  the 
external  dimensions,  in  20  specimens  (measurements  in 
mm):  notum  length  =  29.20,  width  =  13.90.  height  = 
7.14;  width  of  the  sole  =  4.41;  width  of  the  right  h\- 
pouotimi  =  4.14,  distance  from  the  leiiKile  geiiit;il  pore 
fon\artl  =  14.50,  and  behind  =  12.90:  distance  Iroiii  the 
female  genital  pore  to  the  pedal  lurrow  =  0.73.) 


Inlciddl  Addloidij:  Sali\ar\  gland  well  de\i'loped. 
loniii'd  b\  large  and  prominent  acini,  some  (jiiile  loose, 
others  ((Jiiipaet.  .Anterior  iiitestiii;il  loop.  c-o\cred  b\  the 
anterior  lobe  ol  the  digestise  gland. 

The  two  nerxe  pairs,  pedal  ;iiid  palial.  originate  \vy\ 
close  to  the  respectixc  ganglia  ol  the  periesopiiageal  ring 
and  stretch  through  the  ceii(i;il  ca\it\  in  two  sets,  right 
and  left,  which  se|i;iiate  from  e;icli  other  about  a  foiirtii 
ol  the  w,i\  liiiiii  liieii  sl;ir(iiig  phice.  continuing  thus  al- 
most to  the  end  ol  the  cenliai  bod\  ca\il\.  where  thi'V 
c-oiiie  siighlK  closer  and  peneti.ile  llie  inlegmnent  (Fig- 
ure 3).  The  antcaior  aortic  arler\  meets  the  nent's  near 
their  beginning  I  in  ;interior  direction)  (Figure  3).  The 
posterior  pi'dal  biiinih  ol  tin-  aortic  ;irtei-\  was  not 
found.  .\\er;ige  measurements  ol  the  uenes  in  20  spc- 
cimeiits  ill  mm:  tola!  leuglli  -  29.9:  together  =  4.48; 
apart  =  16.5;  maximum  sep;iration  =  2.72  nun;  meeting 
ol  the  aortic  artel^  =   1.76  mm  from  the  origin.' 

Pedal  gland  (Figure  4)  shoil .  Ilalleiied  bro.id  m  ils 
proximal  portion,  lun'rowing  in  the  middle  with  the  pos- 
terior extreiiiit\  rounded  and  somewhat  broadened,  dor- 
sal opening  s(.'allo[)ed:  longitndinalK  three  areas  are  \is- 
il)le.  ,\  tninsliicent  outer  one  on  each  side,  and  a  slightlv 
roiigli  eeiiti'iil  one  that  is  higher,  narrower  ;iiid  \ellowish. 
.\t  the  distal  extreiiiit\  ol  this  central  area  is  the  pene- 
tration scar  ol  till-  inferior  bi;nicli  of  the  aortic  arteiA', 
right  under  the  lu'nc  giingiia.  e\\erage  dimension  of  the 
pedal  gland  in  20  specimens  (in  mini:  length  in  the  ii;it- 
ural  position  =  4.20;  distended  i dorsal  smiace>  =  4.44; 
greatest  width  =   1.90.  ( 

The  thin-tissued  copnl.ilion  bursa  .—  spernuithi'ca  or 
spermatoKtic  gland,  ol  some  aiilhorsl  is  spheroid,  ta- 
pering in  ;i  distill  lengtliening  that  folds  and  joins  the 
bursa  duct  tow;ird  its  base,  receiving  the  junctor  duct  ( = 
canalis  junctor  ol  some  authors i  at  (he  bnrs;i  ti]).  The 
bursa  duct,  ol  more  rigid  tissue,  is  e\lindrical  and  ex- 
treineK  short  in  the  jnxcnile  .ininial.  but  lengthens  in 
the  iiilnlt.  becoming  an  elongated  s|)lieroi(l  bursa.  The 
imietoi  duel  is  lliiek  and  siimons.  luu'rowing  siightiv  in 
its  Hnal  portion,  when,  alter  entwining  part  ol  the  base 
of  the  burs;i  duct,  it  j)euetrates  into  the  nairowest  part 
of  the  copukition  bursa  itself  The  rectum  penetrates  tile 
integmnent  near  the  leinale  genital  pore  (Figures  1.  2). 
No  accessor  gland  :  =  accesson  bursa,  ol  some  authors) 
wiis  found. 

Penis  smooth,  with  a  short,  evlindrieal  \crge.  swelling 
;i  bit  bcMiiid  the  middle  ol  its  lull  length,  forming  on 
the  sides  two  ex|iansions  tluit  iiuirk  tlie  beginning  ol  tlie 
glaus.  (dans  longi'r  than  the  \erge,  becoming  slender  in 
its  distal  portion;  generalK  cuned.  i^etractor  muscli'  of 
the  penis  \er\  \ariable  in  lengtii  (Figures  5-16t.  '{"here 
is  no  sjKitha.  (.\\erage  dimensions  ol  the  penis  in  20 
specimens  tin  mm);  total  length  =  2.26;  greatest  diiun- 
eter  of  the  glaus  =  1.45;  \erge  length  =  1.02;  \erge 
diameter  =   1 .07. ) 

The  sheath  surrounding  the  penis  i  I'"igure  17'  has  two 
distinct  regions.  clearK'  differentiated  b\-  the  well-de- 
fined presenci'  of  two  different  t\pes  of  tissue.  The  prox- 
imal portion  of  tlie  sheatli.  which  surrounds  tlie  penis. 


Paw  82 


THE  NAUTILUS,  \"ol.  116,  No.  3 


prt 


dpp        del    teg 


vao 


Figuivs  1-4.  ilctrrornoiuinii  liiiuii/aim.  1,  2.  I'.ul  (il  llic  postrnor  <j;cnilal  <.p,;uns  anil  nl  llic 
..Itlu'  pedal  ncr\X's  .M(:ra594).  4.  IVdal  Klaiul  (MC:i>6594).  al.c:  opcninsl;  aiir:  iirnv  m.U:  art 
bursa;  dhc;  copulation  hiu'sa  dnct;  del:  junctor  duct;  ddni:  middle  ileferens;  dpd:  distal  poste 
deferens;  glp:  part  of  llie  pedal  iiland;' nnp:  retractor  niiisele  ol  tlie  penis;  npp:  pair  ol  pe( 
oviduct;  pi1:  prostate;  rec:  central  rei;i()n;  rep:  perieardi;il  re',ii()n:  ret:  part  ol  the  reetuin:  te>i: 
branch  oi  tiie  aortic  arteiT. 


leehini  i  \1(  • 
:  anterior  aort 
rior  delerens; 
hil  and  palial 
intei'unieiil; 


(i594i.  3.  .\ 
ic  arten;  hoc 
dpp:  proxiin 
iienes;  (i\  i: 
\;io:  part  ol 


r;nit;enieiit 
copulation 
al  posterior 
part  of  the 
the  inferior 


W.  Tlioiiie  ct  al,  200 


Page  83 


2  mm 


2mm 


Figures  5-16.  llrtcnn  H'^iiiiiui  hiiuiiiniiii  5-13.  'Hiicc  (lilTcriMit  \if\vs  ol' the  penis,  in  -3  adult  specimens  ( respecti\el\-:  MUSM 
2513/h  iFii^ines  5-7).  MU.SM  145S  iFii;ures  S-l()!.  KMNH  594SS/a  (Figures  11-1.3)1.  14-16.  Three  difCerent  \ie\vs  of' the  penis 
in  one  juvenile  specimen  (FMNH  594S6/1)).  dda:  antcnor  poiliou  of  the  deferens  duet:  glp:  ghms:  nnp:  retractor  nniscle  of  the 
penis;  soq;  \erge. 


Pa^e  84 


THE  NAUTILUS.  \"c)l.  116,  No.  3 


2mm 


Hrlrn,va"i,m„l  li„m,,an„    17.  Shcall,  that  r,n,-lops  llir  p<-,ns.  „pr,u.l  Inn.^.t.ulin.dK   '^U^^if^^J^/bl/^- ^'"7 
sheath    sliowinil  the  shaip  l<,nuitucl,H.Ll  folds  ul  thr  iMt.rnal  tis.su.^  (MUSM  2ol;Vh).  19.  Shea  h. 


Figures  17-20 

of  the  ilistal  i)oi'tli>n  (il  the  penis  siieaiii,  sinnviiin  ujc  mi,u|j  iw.i-i,.ni....i.  ,..,,. i        i         i      l  l 

opened  lonf^itndinallv.  sliowing  the  presenee  of  a  sali.Mit  i;l..hnlar  lonnatuH,  nt  du-  intrrn.il  tissue,  similar  t<,  that  desenhed  by 
Hoffmann  (1927-  lor  //.  odhm-ri  as  a  crland  (FMNH  594S(rl.'.  20.  He-un.  ..I  (hr  sah.  „t  .^Inl.nla,  lonnat,,.,,.  ^n-atlN  rnlaru.'.t.  also 
,shoNvine  details  ol  the  tissue  lolds  (FMNII  .59486/1)).  l.ai:  shr.itli:  d.)l):  lolds;  i;ll);  saheiit  i^lohnlar  tonuatu.ii. 


|.  W.  ThoiiK-  (■/  ai.  2002 


Page  85 


is  e\trt'iiiel\'  tliin.  often  allowiii'j;  \isiializati(iii  of  tin-  pv- 
iiis  b\  transparent'.  The  distal  portion.  startin<j;  Ironi  a 
si<4nificant  constriction  ri<j;lit  alter  tlie  tip  of  tlie  penis, 
which  is  swollen,  has  numerons  lon<jitndinal  lokls  inter- 
nally and  a  porous  and  velkjwish  (glandular?)  aspect. 
These  folds  nia\'  lie  more  or  less  tight,  neatK'  arranged, 
and  rectilinear  lespecialK  in  ju\enile  specimens)  or  in 
disarras'  (Figure  IS),  Exen  in  juvenile  specimens  where 
the  male  region  is  little  developed,  the  minisculc  folds 
(Figure  20)  can  be  seen  inside  the  slieath.  The  penial 
gland  is  absent. 

The  mandible  is  formed  bv  an  average  of  2.'3  laths  ( = 
ribs,  of  some  authors),  verv  visible  and  somewhat  sepa- 
rated one  from  the  other.  The  most  central  laths  are  a 
bit  taller  than  the  others.  \o  transverse  grooves  were 
noted,  perhaps  due  to  the  obvious  wear.  The  average 
length  and  greatest  width  of  the  mandible,  in  the  same 
lots  utilized  for  extraction  of  tlie  radulae.  were  2.1  mm 
and  0.5  imn  respectivelv  (Figure  21). 

The  radula  has  a  great  number  of  lateral  teeth,  dis- 
tributed rectilinearlv  in  columns  parallel  to  the  single, 
central  line  of  rachidian  teeth,  which  are  alwavs  smaller. 
An  average  of  10.3  coUnnns  and  120  transverse  rows 
were  counted,  resulting  in  a  calculation  of  12.360  teeth 
per  radula.  The  average  dimensions  of  the  radula  were 
5.85  mm  length  bv  3.41  mm  at  the  greatest  width.  The 
average  dimensions  f)f  20  lateral  teeth  were:  length  56.2 
[xm,  width  24,5  ixm,  distances  from  the  greatest  width 
to  tlie  tip  29.43  |xm.  The  average  dimensions  of  20  rach- 
idian teeth  were:  length  32. S  |i.m,  width  12.8  |i.m,  dis- 
tance from  the  greatest  width  to  the  tip  17.0  p.m.  (Fig- 


ures 


oo_o 


■23). 


NeotApe  (herein  designated):  Ml  S.\l  4001,  from 
tvpe  localitv,  24  |an.  2002,  Icii.  Mina  Ivanii'rc/  and  L'lises 
Zamora. 

^^Jie  Localitv:  Peru,  Dpto.  fJma.  Oerro  San  Cristo- 
bal, ( 12-01 '4.5',  f  \,  77'00'52.6"  W),  215  m  aliove  sea 
level. 

Other  Material  Examined:  .\1LS.\1  4(iOl  neotvpe), 
Peru,  Dpto.  Lima,  Cerro  San  Cristobal  i  12  01 '45.1"  N, 
77°00'52,6"  W),  215  m  above  sea  level,  24  Jan.  2002, 
leg.  Rina  Ranu'rez  and  Ulises  Zamora;  MUSM  4000,  2 
specimens,  same  data  as  the  neot\pe:  MUSM  14.58,  1 
specimen,  Peni,  Dpto.  Lima,  Bcjsque  Zarate  i  ir54'  X, 
76°29'  W),  3000  m  above  sea  level,  24  Mar.  1978.  leg. 
X'alencia  and  Franke:  MUS.V!  1460a,  1  specimen,  Peru, 
Dpto.  Lima,  Bosque  Zarate  (ir54'  N,  76°29'  W) 
(Chourritos  II).  2880  m  above  ,sea  level.  01  Mar,1980, 
leg,  X'alencia  and  Franke;  MUS.M  1660,  4  specimens, 
Peru,  Dpto,  Lima,  Lomas  ck-  .\tf)Congo  (Steinloma) 
(12°13'  N.  76°54'  \V),  28  Sep.  1951,'  leg.  Koepcke 
(Kp340m);  MUSM  18()7a.  3  specimens,  Peru,  Dpto. 
Lima,  Surco  (11°52'  \,  76=28'  W),  leg.  Koepcke 
(KplOlOa);  .MUSM  2388,  1  specimen.  Peru,  Dpto, 
Lima,  Surco  (]r52'  X,  76''28'  W"),  2100  m  above  sea 
level.  17  Aug,  1965;  MUSM  2389,  1  specimen.  Peni, 
Dpto.  Lima.  Distrito  de  San  Borja,  (Rio  Surco),  (I2°06' 


X,  77-01'  W).  08  Juu,  1986,  leg.  A.  Rossi:  MUSM  2513. 
3  specimens.  Pern.  Lima,  Ihiachipa  (12()0'  .X,  76°56' 
W);  .MUSM  3251,  2  specimens.  Pern.  Dpto,  Lima,  Can- 
ta  (Culluhuav)  (iri9'  X,  76°33'  W"),  3550  m  above  sea 
level,  24  Mar.  1989,  leg,  H.  Sisniegas:  SMF  108548  (ho- 
lotvpe  of  Wifiiniiui  {Hi'lcrova^nina)  peruviana).  Peni. 
Dpto.  Lima,  Lomas  (k)  .^tocongo  (i2°13'  X.  7(-)°54'  \V), 
2S  Oct,  1951.  \eo.  Koepcke:  S.MF  32.3293/1,  1  specimen, 
cv-NUSM  40061,  same  data  as  holot\pe;  SMF  140062/ 
7,  7  specimens.  Peru,  Dpto.  Lima,  Cerro  San  (Cristobal. 
leg.  W.  Wevranch;  F.M.XII  59486,  8  specimens.  Pern, 
Dpto.    Lima,   April    1948,   leg.   W.   Wevranch;    F,MXH 

59487,  6  specimens,  Peru,  Dpto,  Lima,  Lomas  do  .\to- 
congo  (12"13'  X,  76°54'  W).  leg.  \V.  Wevrancli;  FMXH, 

59488,  3  specimens,  Peni,  Oconeque,  Puno,  4  Oct.  1941. 
leg.  C.  C.  Sanborn;  FMXH  300744,  1  specimen,  (ex- 
MUS.M  4000),  same  data  of  the  neotxpe:  BMXH 
20020126,  1  specimens.  (e\-MUSM  4000),' same  data  of 
the  neotvpe;  MCP  6501,  1  specimen,  Peni,  Dpto.  Lima, 
Bosque  Zarate.  ill°54"  X,  76°29"  W").  .3000  m  above  sea 
level.  24  Mar.  1978.  leg.  \alencia  and  Franke:  MCP 
6502,  3  specimens,  Peru,  Dpto.  Lima,  Surco  ( 11°52'  X, 
76°28'  \\'),  2100  m  above  sea  level,  17  Aug,  1965;  .MCP 
6504,  1  specimen,  Peru.  Dpto,  Lima,  ("anta  iCullnhnav) 
(iri9'  X,  76°33'  W"),  3550  m  above  sea  level,  24  .Mar. 
1989,  leg,  H.  Sisniegas;  MCP  6593.  1  specimen,  Peni. 
Lima.  Huachipa  (12^00'  X.  76°56'  W);  MCP  6594.  1 
specimen.  Pern,  Dpto,  Lima,  Apr.  1948.  leg.  W.  Wev- 
ranch; MCP  7988.  2  specimens.  (ex-MUSM  4000),  sanie 
data  of  the  neotvpe:  USXM  574941,  1  specimen,  Peni: 
XRS  949(1052!  (holotvpe  of  Vaginiiia  ixlhncril 

DISCUSSlOX 

Vdilinitliis  liiiuii/aitiis  was  brieflv  described  bv  Lesson 
I  1830),  who  used  a  few  moiphological  featiiri's  in  his 
description.  Lesson  also  offered  a  color  plate,  probably 
prepared  in  Peni,  including  dorsal  and  ventral  views  of 
the  animal.  This  specimen  was  not  deposited  at  the  Paris 
Mnseiim,  and  could  not  be  found,  (k'spite  our  efforts, 
in  anv  other  major  ICuropeaii  .Museum.  It  is  vim-%  likeK- 
that  Berard  lost  the  spec  imen  after  the  artwork  was  com- 
pleted. We  conclude  that  there  vv;ls  never  a  holotvpe. 
The  description  and  figures  were  used  bv  Deshayes 
(  1830-18.32)  in  Fenissac  :ind  Desliaves,  with  no  further 
comments. 

.\lth()ngh  the  holotvpe  Vaginirw  (Hetcrocagiiiina)  pe- 
nniaitfi  Krans,  1953,  is  an  immature  specimen,  the  svn- 
onvmvzation  of  this  latter  species  by  Kraus  1 1954)  based 
on  information  from  W.  Wevranch  is  confirmed  alter  its 
examination. 

Recent  collections  ;ind  observations  of  live  specimens 
made  bv  our  collaborator  Rina  Rami're/  confinn  that 
there  is  onlv  (me  species  at  the  tvpe  loc-alitv  and  its  vi- 
cinities. 

Therefore,  we  herein  propose  the  designation  of 
a  neotype  for  the  species  (see  above),  deposited  at  the 
Miiseo  (le  Historia  Xatnral.  Univcrsidad  Xacional  Mavor 
de  San  .Marcos  (MUSM).  Lima.  Peni,  MUSM  4001,  and 


Page  S6 


THE  NAUTILUS.  \  ol.  1  Ifi.  No.  3 


■'■^^'*^^->V 


I  iguii-  21. 


(.>  i„  i,.i  ..,  .V  ,,1  ,,  iM.Muiii.ir  Mcr  :')i.i  1 


Figures  22-23.     Uctciiniiffnina  liiiifii/aiKi:  Mnlillr  region  ol  tljc  ladiihi,  22.  sh^lilK  .ind  2:5.  '.;ri'a(l\  ciil.ii'.^id  .\l(d'  251 
lateral  lootli;  (In  racliidian  toolli  k-ciihal). 


:Vl,i.  d' 


W.  TIioiik'  ,1  a!..  2002 


VdMv  S7 


collecti'd  at  tlu'  nriifiiial  tvpc  Imvilih  dllln'  species.  ( ^ei- 
ro  San  C;rist()l>al. 

Holfiiiann  (1927)  pnipcised  tlu-  species  \'(iiiiitulii 
odhncri,  especialK  due  to  tlie  presence  in  tlie  iienis 
slieath  of  a  lilandular  protuberance.  Tlioine  ( 19S4I  in  liis 
rede.scription  ol  the  holotxpi'  oi  \'  itdhmri.  made  no 
referenci'  to  that  di.stinctix'e  protuherance  liecanse  the 
holotA]ie  was  extensively  daniai^ed. 

In  tlui-e  jn\-enile  specimens  (SWV  10S.54S.  MCF 
6593.  and  FMHN  594Sfi/dl.  amomj;  theni  the  holot\pe 
ol  //.  jicnnidiKi  (  =  H.  liiiuii/tniiiy  an  ahiiost  cncnlai 
protnlieiance  on  the  inner  w.ill  ol  the  penis  sheath  was 
tomnl.  \ei"\  smiilai'  to  that  described  1)\  iloirmann 
(1927)  for  //.  udhiicri  as  a  "'^lantl".  Tlie  ■^land"  de- 
serilied  tor  //.  odhiicii.  used  lo  <hstin<i;nish  it  from  //. 
liiinii/diiii  ("sensn"  Kraus).  appears  to  be  oiil\  a  more 
salient,  globular  lorniation  ol  the  internal  tissue  ol  the 
sheath,  because  its  texture  is  the  same  as  this  tissue  (not 
a  "gland"')  (Figures  19-20). 

Despite  the  additions  antl  changes  uiaile  lien-in.  lh<' 
dichotonions  key  tor  determining  the  .\mericau  geneia. 
proposed  bv  Thome  (1975).  contimies  to  be  valid  lor 
Hcfcroc'dgiitiiia. 

a(;k\o\\ledc:ments 

W'e  are  gratelul  to  William  Beltou,  (uvat  Cacapon,  West 
\'irginia,  U.S.\,  lor  translating  this  article  into  English;  to 
Jose  H.  Leal,  .Sanibel,  Florida.  L'S.X.  loi'  atlditional  help 
widi  the  language;  to  technician  .\ugnsto  Mattos.  of 
PUCRS,  for  the  SEM  photomicrographs:  to  Rina  Ka- 
mirez-Messias,  Coordinator  of  the  Department  of  Mal- 
acolog\-  and  Carcinologv,  Museo  de  Historia  Xatinal. 
Universidad  Nacional  Maxor  de  ,Sau  Marcos,  biiua. 
Peni.  for  collecting,  donating,  and  lending  specimens;  to 
-Margaret  Baki-r  anil  Martin  Pnzdia.  Division  ol  Inver- 
tebrates. The  Field  Museimi  of  Natural  Histon,  Chica- 
go, Illinois,  USA;  and  to  Ronald  [anssen.  Curator  ol  the 
Mollusea  Section,  Farsehimgsinstitut  und  Naturnmseinn 
Senckenberg,  Frankfnrt-ani-Main,  (lernianv  tor  the  loan 
and  donation  of  specimens.  Work  prepared  with  finan- 
cial support  from  CNP<i,  CAPES,  FAPEHCS  and 
PUCRS  (Brazil). 

LITER.4TURE  CITED 

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29-46, 

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THE  NAUTILUS  1  lfi(.",):S9-94.  200 


Page  89 


Aiiatoniy  of  the  dromedaiy  jiiniping-slng,  Hciiij)hilli(i 
dwmcdanus  Branson,  1972  (Gastropoda:  St)l()nnnat()plio)-a: 
Arionidae),  with  new  distributional  records 


Kristiina  ()>aska 

Hidliiix  EnxiroiiiiHiit.il  Hisiarcli  lid, 

4 ISO  Clinton  V\mv 

X'ietoria,  British  (  Nilnniliia 

CANADA  \'SZ  (SMI 

and 

Department  of  Forest  Seicincs 

Universit\  nl  liritisli  (jiliiinlna 

Vaneoincr,  Britisli  C^oliniihia 

CANADA  V6T  1Z4 

kcnaskaCn'jdniicni.eoni 


Lylf  C'liiclu'sliT 

2S()5  (Jreenhriar  Uonlexard 
Wellington.  FL33414  USA 


Ileike  Reisi- 

State  Miiseniii  ol  Natural  ilistoiA- 
PF3()0  ir,i 
D-()2S0(i  Coilit/ 
CKHMANY 


\\illiain  P.  Leonard 

223  Foote  Street  NW 
OKnipia.  \\A  9S5()2  USA 


Jim  Haiii^li 

2()|S  Dn  Creek  Hoad 
Flleiislmp'   W  \  9SV)2(i  USA 


ABSTRACT 

rile  '^eniis  Hiiiijiliilliii  I'epicseiited  1 1\  al  least  7  spi-rii-s,  is  a 
poiiilv  kiiiiwii  '4riiiip  III  sjii'^s  I'lidiiiiie  Id  wcslirii  \im1Ii  \nier- 
iia  We  niNcsti^atid  tlic  disl nl iiilii m  ,uid  liahitats  ol  //  dni- 
iiiiddiiiis  Branson.  1972.  and  present  a  redescription  ol  tlie 
anatoins  ol  its  distal  genitalia,  nsetl  tor  speeies  identiliiation. 
We  collected  specimens  from  fi  localities  on  X'aneoiner  Island, 
British  Columliia,  Canada,  and  hom  9  localities  in  Wasinngton 
State.  USA.  The  \'ancon\er  Island  localities  repnsmt  a  north- 
ward range  extension  loi'  the  species  and  first  confirmed  re- 
cords Iroin  ("anaiki  rlii-  species  occurred  from  ncLir  sea  le\el 
to  an  eliAatioii  ol  I  :i70  in.  wilh  iiiosi  localities  alioxe  701)  nl. 
In  all  disscctt-tl  specimens  lat  least  I  per  local it\!  the  penis  was 
priiximallv  broad  and  distalK  taperetl;  a  \\(ll-ile\ eloped  \erge 
was  present  witliin  the  penis  sac.  Contran  to  the  original  spe- 
cies description,  the  penis  had  an  accesson  sac  that  inserted 
distalK  near  the  gonopore,  .Although  the  anatonu  of  the  gen- 
italia did  not  match  the  original  description,  the  dissection  ol 
the  t\pe  material  confirmed  that  onr  speiiineiis  repnsented  // 
ilntiiiiildriii.s. 


IXTHODUCTION 

The  genus  UcuipJiiUid  lilaiul  and  Hniiie\.  1  S72  eiinsksts 
1)1  a  giDiip  <il  arloiiici  slugs  endeinic  In  weslein  North 
America.  Slug.s  of  thi.s  genus  are  cliaiaelen/ed  li\  a  vis- 
cera! ca\ih'  ele\ated  into  a  pniiiciiinced.  dorsal  liiiinp.  a 
.shell-plate  that  i.s  partialK  exposed  llirongh  a  slil  in  liie 
mantle  (Pilshn.  1948).  and  a  remarkable  anti-|)redatoi-\ 
hehaxior:    the\    txpicalK    exhibit   "xioleiit   writhing   and 


leaping  in  response  to  iiiolesladoii  I'ilslirv.  I94S;  73S). 
Sex'eii  species  are  eiiireniK  reeogni/ed  //  'J/liidlilostl 
Rlaiid  and  Hiiniev,  1S72:  //  hnrrhfjoiii  I'ilsbrx.  1948: 
//,  pinillirmiii  lii.inson  197")  //  idiiichis  l'ilsbi"\  and 
N'anatta.  1897:  //  ilriiinitltiiiiis  liianson,  1972:  //.  daii- 
iiisi  X'aiiatta.  191  I:  and  //,  iiiulonri  Pilsbn.  1917:  Tiir- 
geon  et.  al.,  I998i.  i)iit  :iddilli)ii:il.  midescriiied  species 
ma\  exist  i  kelle\  el  al,.  I999v  I'lie  geims  has  receixed 
little  attention  lioiii  ell  In  a'  s\  slenialists  or  ecoiogists.  and 
file  taxononiie  nkitionsliips.  disliibnlion.  and  ecologx  ol 
all  speeies  ;iic  poorK  known. 

Hr;nison  11972'  described  //  drdmidiiriiis  based  on 
specimens  Ifom  the  OKnipie  Nh)milains.  Washington. 
His  di'seription  was  based  primariK  on  external  chaiae- 
leiislics  w  illi  lillli'  rerer<'nce  lo  llie  anatonix'  of  the  ilistal 
reprodiieli\e  s\sleiii.  wliieh  is  ollen  extremelv  useful  or 
e\fii  essential  loi' the  idenlilicatioii  ol  shigs  (Keniex  and 
Cameron,  1979:  dompa.  1984:  (;onie/,.  20()Ll  Onr  in- 
leicsl  ill  llns  speeies  arose  alter  the  diseovcn  in  1999  ol 
ii  large  ((iO  iiiiii  In  length;  jnmpiiig-shig  Ironi  N'aneouxcr 
Isl.uid.  Hnlisli  (  ohmibia.  (Canada:  onK  //.  '^laiidulosa.  a 
iiiiieli  sni.illc'r  speeies.  was  pre\ionsl\-  known  Iroin  tiie 
west  coast  of  liritisli  (.'ohunbia  i  Pilsbrx  I94S:  Cameron 
198fSi,  but  an  earK  report  of  a  huge,  unidentified //(7»- 
phillia  loriginalK  niisidentified  as  //.  inaUmei)  existetl 
from  \ancou\cr  Island  (Ihinham.  192fi'.  Our  attempts 
to  \erif\  the  identitx  of  the  new  s])e(,'imen  were  Irus- 
Ir.iled  l)\  ,1  lack  ol  piiblisiied  information  on  the  genitalia 
of  //  di(iiii(d<inus.  iM'omjiting  us  to  examine  the  t\pe 
mati'rial  and  to  collect  additional  specinu'us  from  Wasli- 


Pat:.'  yo 


THE  \AUTI1AS,  \nl.  lift.  \< 


iiiiitoii  State.  United  States,  lor  eoinpaiisoiis.  Here  we 
present  a  description  of  tlie  distal  reproductixe  anatonu 
of//.  (Iroiiu'daiiits  based  on  tliesc  findings.  We  also  \ivc- 
sent  new  distribntional  records  tor  the  species,  inclndiiig 
a  northward  range  extension  and  the  first  confirmed  doc- 
niiientation  of  the  species  from  ( 'anada. 

MATERIALS  AND  METHODS 

We  colleetetl  slugs  from  forestetl  areas  on  Nancouvt'r  Is- 
land. British  (]olmnl)ia.  (Canada,  and  in  Washington 
State.  United  States,  between  1999  and  2002  (.seeAp- 
pendi\  loi'  collection  localities  antl  tlatcs).  On  X'ancouver 
Island,  the  first  specimen  of  H.  (Iroiiicihiriiis  was  found 
incidentalK  during  sunevs  for  terrestrial  gastropods  in 
the  autumn  ol  1999.  but  sunexs  in  snbseijnent  vears 
focused  specificalK'  on  potential  habitats  tor  this  species 
(a  total  ol  104  forested  sites  were  suncxed  on  \'ancon\er 
Island  and  3S  sites  on  the  mainland  of  British  Oolnm- 
bia).  Specimens  from  Washington  wfre  colleetetl  either 
for  comparisons  with  material  trom  \'ancou\er  Island  or 
during  opportunistic  suneys  in  \arions  localities.  We  lo- 
cated slugs  b\  turning  over  woocK  debris  and  b\'  xisualK 
searching  other  potential  microhabitats  on  the  torest 
floor.  Specimens  collected  were  killed  b\  drowning  anil 
preser\^ed  in  either  10%  ethanol  or  isopropxl  alcohol, 
and  vouchers  were  ileposited  in  the  eolk'ction  at  the 
Ho\al  British  Oolumbia  Musemn  (RBOMl.  Mctoria. 
British  C^olnuibia.  (.'anada. 

We  dissected  at  least  one  specimen  from  each  localit\'. 
The  dissection  ol  new  material  was  pertormed  b\'  LC 
(most  specimens)  and  KO  under  7-30X  magnification 
using  a  dissecting  microscope.  Drawings  of  the  repro- 
ductixe sNstem  of  selected  sjiei'imens  were  made  using 
camera  lueida  attached  to  a  (.lissecting  microscope.  The 
dissection  and  examination  of  the  holotxpe  (USNM 
577690;  National  Museum  of  Natural  Histon,  Washing- 
ton, DO)  and  txvo  paratxpes  (FMNH  173022',  Field  Mii- 
.semn  of  Natural  llistorv.  Chicago;  DMNH  43029.  Del- 
aware .Museum  of  Natural  llislon.  Wilmington)  was 
performed  by  HH. 

RESULTS 

On  Nanconxcr  Islaml  we  located  //  (Inxncddiiiis  al  six 
localities  ranging  from  ni'ar  sea  le\cl  lo  ;iu  elevation  ol 
1060  m  :.\j)pen(li\).  These  loc-alities  represent  ,i  north- 
ward range  extension  for  tiie  species  (  Eignre  I  1.  I  iabilats 
consisted  ol  conilerous  forest,  dominated  !)n  western 
hemlock  [Tsnou  liclrrophijlla).  western  redeetlar  {'ihuja 
pUcata).  and  am.ibilis  fir  (Ahics  aniahili.s)-  the  forest  tloor 
wa.s  txpically  composed  of  thin,  compact  mcdle  littei 
Fixe  of  the  six  loe;ilities  xxere  in  ri'nmant  patches  ol  Old 
groxxth  forest  (with  trees  >  250  years  old);  one  loc;ilit\ 
(record  3  in  .\ppendix)  was  in  second  growth  torest  with 
some  old  trees.  All  sites  contained  abimdant  coarse 
woody  deliris,  inclnding  large-diameter  logs  in  atlxanced 
.stages  of  (k-cay.  In  Washington,  habitats  of  H.  drouw- 
dariuH  consisted  of  both  old-groxx'th  and  older  second- 


growth  conilerous  forests  dominated  bx  Douglas-fir 
[Fsvudotsiiiid  iiwiizicsii)  and  xxcstern  hemlock;  mountain 
hemlock  (7's7(g«  mciicusidtKi^  anil  subalpine  fir  iAhii's 
hisiocarpa)  xvere  present  at  the  highest  elexation  sites. 
Within  these  habitats,  xxe  tonnd  //.  drotiicddriiis  under 
and  xxithin  decaxing  logs  and  lallen  bark,  under  rocks, 
and.  once,  actixe  on  the  surface  of  the  litter.  Most  sites 
xxere  mesic,  but  at  three  high-elexation  sites  on  the  east 
slope  of  the  Cascade  Mountains  (records  10,  11.  and  13 
in  Appendix  1)  the  slugs  occurred  on  drier  talus  sub- 
strates. 


.Moiii'iioi.oi.x   wii  Intkhnai,  .AxxToxiv 

Extenuillx,  all  specimens  xxere  similai'  and  contormed 
xxith  the  tiescriptioii  b\  Branson  (1972)  (Figure  2).  The 
pretlominaut  color  ol  the  mantle  and  foot  xxas  various 
shades  ol  grax  xxith  cream-colored  mottling  on  the  sides; 
the  sole  of  the  loot  xxas  pale-xelloxx;  orange-xelloxx'.  or 
cream.  Otic  inilixidual  (Irom  loealitx  1  in  Appendi.x)  xxas 
bull  broxxn  xxith  a  bright  xelloxx  sole.  The  tail-portion  ol 
the  foot  xvas  laterallv  compressed,  keeled,  and  tipped 
\xith  a  horn-like  protuberance  leaudal  horn),  as  de- 
scribed bv  Bran.son  (Figure  2.  insert).  The  nine  \'aneon- 
xer  Island  specimens  ranged  from  40  to  60  nun  in 
length;  three  specimens  trom  Washington  (record  9  in 
.Appendix)  xx'ere  32-37  unn  (all  measurements  trom 
fresh,  reliLxed  specimens  after  droxxuing). 

Inteniiillv.  the  oxotestis  xxas  grax-broxxn  antl  similar  in 
color  to  the  digestixe  gland,  ("lose  to  tlie  entrance  ol  the 
ducts  ol  the  digestixe  gland  there  xxas  an  intestinal  di- 
xerticnlnm  ol  nnknoxxii  hmction.  The  penis  xxas  proxi- 
inallx  bnnul  and  tapering  distallx'.  and  a  xxell-developed 
xerge  x\as  present  (Figure  3).  The  xerge  filled  most  of 
the  broad  end  ol  the  penis  sac;  there  xxere  no  other 
intra-penial  structures.  The  penial  retractor  nmsele  in- 
serteil  at  the  long  and  slender  epipluillus.  The  penis  had 
an  iiccesson  s;ie.  xxliicli  eonsist("d  of  ;i  narroxx  tube  that 
broadened  pi'oxiuKillx  and  iiregularix  i  Figure  3!.  The  tu- 
bular base  of  the  accesson  sac  inserted  toxxards  the  dis- 
tal end  ol  (he  penis  ne;u'  the  gonopore.  The  bursa  I'o- 
[iilkitrix  (spermatheeal  consisted  ol  a  stout  tube  (h;il 
broadened  proxiuuilK  into  a  globular  sac.  more  jiio- 
nonuced  in  some  iudixidnals  than  in  others.  The  peins, 
Iree  oxiduel,  anil  liiusa  eopillatrix  joined  close  to  the 
gonopore;  iheie  \\,is  no  distinct  alrinm. 

The  holotxpe  is  a  lullx  iidnll  specimen  with  ;i  Lirge 
.illinmen  ghmd.  ;ind  its  genital  aiuitomx  is  generalK  the 
s;ime  as  m  oni  siiecimens.  The  distal  part  ol  the  penis 
is  liilhei-  u;iii"o\x,  ,ind  the  proxim.il  p.irt  is  wide.  There  is 
a  penial  ;i|)peudi\  that  insi'rts  dist;illx  near  the  gonopore. 
■file  retr;i(lo|-  inserts  .it  the  <'piph;illus.  Intern;ll  penis 
stnietures  xxere  not  inxestigated  to  minimi/.e  damage  to 
llii'  lx|)i'  Sjieeimeu.  Both  |)aratxpes  are  juxenile.  ;ukI 
their  genitalia  are  in  somexxhat  miu,itm:il  positions, 
which  furthei'  hinders  the  inxestigation  (parts  ol  the  dis- 
l.il  genitaliii  ol  speeiiiieii  DMNH  43029  are  xxithin  the 
right  leutacle.  and  those  ol  specimen  F.MNH  173022  ;ire 
partlx  exerted  through  the  genital  pore — both  probablx 


K-  ()\aska  i1  al.  2002 


I'a.'c  91 


Figure  1.       Distil  I  iiitidii  map  lor  lotalitx  iccdnls  ol  U(iiii>hilliii  (Inmnddniis   Solid  t-ircles:  this  si  ink  i  iiiiiiilicis  {oncspoiid  to  those 
ill  AppiiulK);  open  circles:  piv\iiiiis  Iciealltles  (Branson,  1972.   1977,  19S()).  Arrow  points  to  the  t\pe  loealitv  iHrarisoii.  1972). 


artifact.s  of  killing).  Nexertiiele.ss,  a  penial  appendix  i,s 
pre.seiit  at  lea.st  in  specimen  DMNC  43029,  and  the 
stnicture  of  the  everted  genital  complex  in  s|)('ciiiien 
FMNH  17.3022  suggests  its  presence.  \\\  lliicc  t\pc 
specimens  ha\e  the  intestinal  dl\(■l■|i^■llllnll  inciiliinicd 
above. 

DISCUSSION 

The  geographic  tlisti'ihntion  ol  //.  (Ironicilnniis.  like  thai 
of  man\  other  terrestrial  gastropods  in  western  North 
.America,  is  poorK  knowii.  Pre\ious  piiMished  loealit^  re- 
cords aic  iioiii  the  Cascade  Range  and  ()l\in|)ic  I'eiiiii- 
snla  in  western  Washington,  including  the  ()l\iiipic  and 
Mt.  liainier  National  Parks  ami  the  .\lt.  Baker  and  Suo- 
(juahnie  National  Forests  (Branson,  1972.  1977,  19S(),i 
(Figure  1).  In  addition,  some  ol  the  records  Iroiii  west- 
ern Washington  reporteil  1)\  i'ilsliiA  (i94Si  lor  //,  inn- 
loiui.  prior  to  tlie  description  ol  //.  (Iroini-iliiriiis.  may 
lielong  to  the  latter  species.  Our  records  Iroin  Nancouxcr 


Island  represent  a  nortliuard  range  extension  and  first 
c-onfirmed  records  from  Canada.  On  the  mainland,  both 
the  iiortliern  anil  southern  extent  ol  the  species'  distri- 
bution remain  poorK  known.  The  norfliermnost  main- 
lanil  n'cord  is  Iriiiii  the  Cascade  .Mountains.  Wasiiington 
(SiKcr  I'ir  Camp.  .Mount  i5aker  National  I^'orest:  Bran- 
son, 19S0).  close  to  tile  Canadian  border  (  Figure  1  ).  The 
sontherimiost  confirmed  localit\'  is  from  the  (iillord  Bin- 
chot  National  i'"orcst  (record  S  in  .\|')]iendix).  lliiiijiliilliti 
thai  arc  similar  in  si/e  and  external  appearance  to  // 
(Iroiiiciliirius  lune  been  iomid  lartiier  south  in  the  (Cas- 
cade and  C^oast  ranges  ol  northwestern  Oregon  (Tom 
Biirke.  pers.  conmi.  b\  WBI,:  Carol  i^icklord,  pers. 
coiiim.  b\  Wl'I/i.  but  iiecause  these  were  not  dissected 
sjiecics  i(k'ntificatiou  remains  uncertain. 

We  fonnd  H  (Iniiiu-tltirius  at  six  localities  on  X'ancou- 
\cr  Island.  \  prexioiis  report  (llaiiham,  1926:  143)  exists 
of  tx\-o  specimens  of  a  large,  black  jmnping-slug  from 

.  .  the  iioriler  ol  a  good-sized  lake  |most  likely  Ifolyoak 
Lake!  on  Mt.  Brenton.  \'ancou\cr  Island,  at  an  elevation 


Page  92 


THE  NAUTILUS.  \ol.  llfi,  NO.  3 


Figure  2.  Specimen  ot  Hciiiphilliii  droinfdarius  Irniii  Loss 
Oreek.  N'iiiieonver  Island,  Britisli  Colnmhia.  C^anada  (l()calit\'2 
in  Appendix;  KBCM  ()()1-0()2S(M)1)I );  candal  liDin  (.1  llic  same 
animal  linseitl.  Tlu'  Irni'th  ol  tin-  li\c'  animal  was  55  imii. 


Iiitween  3500  and  4000  ft".  The  slugs  were  identified  as 
//  iiifiloiu'i  In  Walter  |.  Excrdani  (of  .Seattle.  Wasliing- 
(i)ni.  apparentK  nn  the  hasis  ol  Ilaiiliani's  later  descrip- 
tion oi  their  color;  no  spi-ciinens  were  a\ailal)le.  as  their 
soft  anatomy  had  disintegrati'd  before  the  slugs  coukl  be 
preserved  (Hanham,  1926).  Later  authors  (Pilsbn',  1948; 
Kozlolt  and  \'ance,  1958)  questioned  the  identification 
ol  this  rcioicl.  which  has  remained  enigmatic.  We  con- 
firmed the  presence  of  H.  clmnwdariiis  near  Holvoak 
Lake  I  record  4  in  .Appendix);  most  likelw  the  slugs  re- 
ported b\  lianliain  Ironi  tlie  same  localitx'  75  vears  pre- 
\iousK  were  also  ot  this  species. 

Our  localit\'  records  trom  \'ancnn\ei'  Island  were  Iroiii 
older  torests  at  elevations  lioni  near  sea  le\cl  to  lOfiO 
m.  A\\  three  localities  in  the  drier  southeastern  and 
soutli-central  parts  ol  the  island  were  abo\'e  TOO  ni  in 
elexation.  and  the  species  seemed  to  be  absent  from 
lower  slopes  ;ind  \alle\'  bottoms.  In  contrast,  the  three 
localities  on  the  wetter,  west  coast  ot  the  island  were 
near  sea-level,  in  Washington,  pre\ious  Records  for  the 
species  existed   tiom   elevations  of   23S   m   to    1436   m 


1  mm 


Figure  3.  HcpnuliictlM-  system  oi  llcini}liilliii  (Inmiiilfiiim  drawn  iroiii  1  spciiniciis  Ironi  Wliilc  Pass.  Lewis  Co..  Wasliiiiuldii 
(HB(,M  0()1-2HB-()()I:  locality  S  in  .\ppendixi;  genitalia  of  specimen  1  and  lateral  \iew  of  the  verge,  dissected  from  tile  peiiial  sae 
of  specimen  2  (insert  I.  .■VC:  aecesson-  sae  of  penis:  .\(.',:  alhnmen  tjland:  BC:  bursa  eopniatrix  ispermatheea);  EP:  epiph:illns;  III); 

hcrmapliroditie  (hiet;  OT;  ovotestis;  V    penis:  I'W:  pcnial  v\all.  ]i(c'led  liaef:  HI   pcni;il  retnietor;  Sl'0\" — sper xulud:  \';  verge; 

\'S:  Vits  ileferens. 


K.  Ovaskar/  ,;/..  2002 


I'asic  9.) 


(Branson.  1972.  19S01.  .Ml  hut  one  iifiiiir  n'conls  Ironi 
W'asilington  (ivconl  7  in  .\ppt'iuli\)  were  lioiri  rclatixcK 
higli  elexation.s  (744-1370  ni).  On  the  east  slope  of  tlie 
Cascade  Hant^e  (records  10-15)  the  species  appeals  to 
be  confinet!  to  woodeil  nioinitaiii  peaks  amidst  drier 
grass/shrul)  hahitats.  Iiiterestiinj;l\.  at  tiiese  sites  the  slugs 
inhabited  sparseK'  wooded  siibalpine  tains. 

E.xternalK-  onr  specimens  conlormed  with  tin-  descrip- 
tion for  //,  (Inniuilarius  (Branson.  1972'.  aIthoir.^h  oin 
measnreineiits  ol  the  length  of  Iresh  specimens  Iroiii 
\ancon\fr  Island  i40-tS()  mm'  were  considi'rabK' greater 
than  the  length  (24-31  mm  I  reported  In  Branson.  .\s 
the  t\\(i  juNcnile  parat\pes  suggest,  most  ol  Bransons 
specimens  might  not  ha\t'  been  hilK  grown.  .Xlthoii'^h 
tlie  anatonn'  ol  the  genitalia  ol  our  specimens  difleied 
from  the  original  description,  thest-  specimens  were 
comparabh'  with  Branson's  il972i  t\pe  material.  Tiie 
original  di'scription  referred  to  the  anatomv  of  the  gen- 
italia three  times:  in  the  idetitilication  ke\  (p.  103):  "pe- 
nis broad,  lacking  an  accesson  gland";  in  the  corrobo- 
rati\e  di'Scription  (p.  105):  "the  inllated  penis  does  not 
bear  an  accesson  sac":  anil  in  the  diagnosis  (p.  106):  "It 
[H.  clniincddriii.s]  differs  from  //,  (laiiiclsi  in  matters  ol 
coloration  and  in  possessing  the  inllatetl  penis  and  in 
lacking  an  accessoiT  stinnilator".  We  presume  that  the 
accesson-  organ  in  question  refers  to  the  same  stnictnre. 
In  the  ailiilt  spei'imens  iinestigated  b\  iis.  onK  the  prox- 
imal portion  ol  the  penis  was  broati,  whereas  its  base 
was  long  and  rather  narrow.  Also  contran'  to  thesi'  state- 
ments, ill  the  specimens  we  examined,  including  t\j)v 
specimens,  a  penial  accesson  sac  was  alwaxs  present. 
However,  the  liolotxpe  as  well  as  the-  two  imestigated 
paratxpes  hail  not  been  disseited  In  Branson,  so  it  re- 
mains unknown  wiiat  specimen,  and  therefore  species. 
he  might  ha\e  cxaiiiiiK-d  lor  his  anatomical  descriptions. 

In  H.  (Iroiiu'claiius.  there  were  no  intrapenial  acces- 
son' stnictures,  such  as  the  "stimulator"  described  lor  H. 
Iiuiloiwi  (Ko/lorr  and  \aiicc.  195SI.  .\  more  detailed, 
comparative  inxi'Stigation  ol  the  xcr^e  and  other  iiitr;i- 
penial  structures  among  species  ol  I Iciiijiliilliii  awaits  in- 
\estigatioii. 

Three  other  larij;e-liodii-d  (>.30  mm  in  Ic-ngth  i  sjieiies 
ol  Hciiijihilliii  are  currently  recognized  iH.  nuilmici.  II 
caiuclus-  and  //.  diinielsi).  A  penial  accesson  sac  is  ab- 
sent in  //  inalonci  (Pilsbn'.  194S;  LC".  unpublished), 
whicli  is  sxnipatiic  with  //  ilroiindiirins  in  the  OKnipic 
Mountains  ol  Washington.  I'AternalK.  the  tx«)  species 
can  be  distinguished  In  the  absence  ol  a  caudal  hum 
and  the  presence  of  a  prominent,  light-colored,  mid-dor- 
sal stripe  on  the  tail  in  H.  iiKihinci.  Hcmpliillin  rniiitliis 
from  eastern  l^ritish  Columbia  and  Washington  also 
lacks  both  a  caudal  horn  and  a  penial  accesson  sac  (Pils- 
bn', 1948).  Of  the  described  species,  H.  droiucdarius 
most  cIoseK  resembles  //.  dtni'uhi^  which  occms  in 
Montana  (Pilsbn',  194S).  ami  our  speciniciis  ke\c(l  out 
to  this  species  using  Branson's  (1972)  identification  key. 
Sx'steinatic  relationships  within  Urinpliillia  and  among 
related  genera  ha\e  not  been  examined  recentK,  and 


comparative  studies  ol  reproductive  analonix  and  genet- 
ics ol  Hiiiijiliilliii  are  in  ordia'. 

ACKNOW  II'DCMIA  rs 

I  he  first  //,  di'oiiicddriiis  Irom  \aiicouver  Island  was 
lonnd  diirinii  gastropod  snnevs  sponsored  bv  W'evcr- 
haeiiser  Canada  (  N'anaimo  Office*.  .Subsccjiienl  financial 
support  lor  sunevs  in  British  (.'olnmbia  came  from  the 
Endangered  Sjiccics  Hecoven  fund.  Wildlife  Habitat 
Canada.  :uid  Dipartmcnt  ol  forest  Sciences.  L'ni\ersit\ 
ol  British  (!olimiliia  ithrouiih  a  Forest  Bencwal  Hrilisli 
(^olnmbia  grant  to  |ohn  Hichardson!. 

We  thank  Su/annc  Beaiichesne.  Christian  Engelstoft. 
Hobert  l-'orsvth.  Brent  lladdawav.  Lee.Ami  Hancock. 
Kellev  joigcnseii.  'rum  Kogiil.  Brad  Moon,  Bill  Null,  ami 
L.emiart  Sopnck  lor  entliusiastic  help  with  the  collection 
of  speciiiR'HS.  Bobert  Forsvth  provided  literature  refer- 
ences and  contacts.  Clen  Dunsworth  and  David  Lindsav 
]ii'o\ided  access  to  forestn  lands.  KelK  Sendall  and  Phil 
Lamiiert  allowinl  us  to  use  the  tacilities  at  the  Boval 
British  Columbia  Museum  and  acce|)ted  our  specimens. 
We  are  also  grateful  to  Hobert  llershler  Timothv  Pearce. 
and  |ii(licii  (icilier  lor  the  loan  ol  tvpe  specimiMis.  Ira 
W  illev   hclpiil  with  the  iiiKi^inn  ol  tlie  figures, 

LITEB.\TUHK  (  ITEl) 

Bransdii.  B.  .\  li)72  lU  iiiplitllid  ilioincdiiriiis.  a  new  ariouiil 
slu'4  Ik  III!  \\asliin>j;t<iii.  'I'lic  Xaiililiis  S5:  100-106. 

Branson.  H,  .\.  I!)77.  I'rcsliwatcr  ami  terrestrial  .Mollusca  ol 
the  Olv  iii|iK'  I'ciiiiiMila.  Wasliiiiiildii.  'I'lic  N'cli^er  U):  319- 
330. 

Biaiisoii.  B,  \.  I9S0.  (.■()llccti()iis  ol  ijastropiKis  Iroiii  the  C'as- 
(■;i(lc  Mduiitaiiis  ol  W'asliiiimon.  'I'lic  N'eliiler  23:  171-17(1 

(ioiiuv.  H  I  2()(ll  Slnirture  aiiilliiuclioiiing  1)1  the  reproiluc- 
tivc  svslciii  In  B.irkcr  <.'■  \1  c'd.!.  The  Biolos^ol  Ter- 
restrial M.illiiMs  (  \BI  ruMi-.liiiiu,  Wallinuford.pp.  307- 
330 

llaiiliani.  .\  W  1020.  Ih  iiiphillui  iiuilnini  \'aii.  The  N'.iutihis 
39:  143-1  It, 

K.llev.  H.,  S.  D.mhiii.  \.  Duncan  and  T  Biirke.  1999.  Pield 
Guide  to  Suncv  and  .Maiiaije  Terrestrial  Moliusk  Spi'cies 
from  the  N'ortlivvest  Forest  I'laii.  Bureau  ol  Laud  Man- 
auinicnl,  Orenoii  Slate  Office.   114  pp. 

Kiincv  M  I"  and  B.  .\.  D.  Caiiierou.  1979.  A  Fiekl  V,\m\c  to 
land  Snails  of  Britiiiu  and  Nortli-we.st  Europe.  William 
(;()lliiis  and  Sons.  Ltd..  Foudoii,  2S8  pp. 

Kozloff  F..  \.  and  1  \ancr.  I95S.  Svsleniatic  status  oi  llcin- 
phiU'ui  iiiiili'iiri    the  Nanlilus  72:  42—19. 

Bilshi-i.  II  A,  19IS.  I.,ind  Mollusca  ol  North  .America  i north 
of  Mexico'.  The  .Acadeinv  ol  Natural  Sciences  ol  I'liila- 
delpliia.  Mononrapli  2:  521  - 1 1  F5.  i-xlvii. 

4'iiinpa.  A  S  I5)SI  I  .and  snails  iStvlommatopliora'.  In:  Toiiipa. 
.\.  S..  N.  II.  X'erdonk  and  j.  .\.  M.  van  den  Biggelaar 
(eds.).  Tlie  .VIollusca,  7:  Beproduction,  .Xcademic  Press, 
Fondoii  ;nid  New  York.  pp.   17-140. 

riip.icoii.  I)  1)  I  I'  (,)uinn  jr  F.  \  Co.ni.  F  (.'■  Hochherj;. 
W.  (;.  Iaoiis^  I'  M  Mikkilsen.  B.  J.  Neves.  C.  F  E.  Boper 
(;.  Boseuherii.  B  Both.  \.  Selieltema.  F.  G.  Thompson. 
M.  X'ecchioue.  and  j.  1).  Williams.  199S.  Common  and 
Scientific  Names  oi  .Xciuatic  luvertehrates  from  the  Fnitcs 


Page  94 


THE  NAITIIA  S.  \  ol.  IKS,  \o.  3 


States  and  Canada:  Mollusks.  Secfind  edition.  American 
Fisheries  Society  Special  Publication  26.  526  pp. 

APPENDLX 

Lxjcalities  for  Hemphillia  (Iroinedarius  examined  for  this 

stiid\'. 

Vancouver  Island.  British  Columbia.  Canada: 

1.  5  km  northeast  of  ^^'ild  Deer  Lake,  9  km  soudi- 
west  of  Duncan  (logging  road  LIOOO):  elexation 
700  m  above  sea  \e\e\  (asD:  4S°41'  N.  123°46'  W: 
S  October  1999  (collected  b\  K.  0\aska.  L.  So- 
piick.  ;uid  S.  Beauchesne):  RBCM  001-00279- 
001. 

2.  Loss  Creek,  at  Www  14  southeast  of  Port  Ren- 
frew; 48°28'  X.  124H.5'  \\';  elevation  49  m  asl:  14 
April  2000:  (collected  b\  R.  Fors\th  and  K.  0\as- 
ka):  RBCM  000-00145^)01:  lo'june  2001  (col- 
lected bv  K.  0\aska  and  C.  Engelstoft);  RBCM 
001-(X)2'S0-001  (Figure  2). 

3.  Banifiekl  West:  4S=.5()'  X,  125°8'  W:  eIe\adon  < 
20  m  asl;  23  JuK'  2000  (collected  b>-  K.  0\;xska: 
maintained  in  capti\ih  until  19  December  2000): 
RBCM  001-00281-001. 

4.  \'anc-ouver  Island,  Mt.  Brenton  (south  end  of  Holv- 
oak  Lake),  ca.  8  km  southwest  of  Chemainus: 
48°.54'  X,  123°50'  \\^;  elevation  1060  m  asl;  25 
.August  2001  (2  .specimens  collected  b\'  K.  0\aska 
L.  Sopuck.  and  S.  Beauchesne);  RBCM  001- 
002S2-001. 

5.  Mt.  Hooper  (shores  of  an  unnamed  lake),  ca.  20 
km  northwest  of  Youbou;  48=^60'  X,  124°29'  ^\"; 
eIe\ation  850  m  ;isl;  10  September  2001  (2  spec- 
imens collected  bv  L.  Sopuck,  C.  Engelstoft,  K. 
Ovaska,  and  S.  Beauchesne);  RBC.VI  001-0028.3- 
001. 

6.  Indian  Creek,  ca.  9  km  X  of  Ucluelet  on  KemiecK 
Flats:  49°02'  X.  125=32'  W":  ele\ation  <  20  nv.  18 
Xo\ ember  2001  (collected  by  L.  Sopuck,  B.  Beas- 
ley,  K.  0\aska,  W.  Leonard,  G.  Shreiber,  and  J. 
Ziegltnim)  (specimen  used  for  genetic  studies: 
photograph  in  persoiuil  files  of  KO). 


\Va.'ihin0on  State.  United  States: 

7.  Clallam  Co..  4.8  km  south  of  BKii  (off  Woods 
Road),  Olvmpic  Xational  Forest;  47°59'  X, 
123°00'  W":  elexation  150  m  asl;  20  Ma\-  2001  (col- 
lected b\  W.  Leonard  and  B.  Moon  >:  RBCM  001- 
00285-001. 

8.  Lewis  Co.,  Gifford  Pinchot  XF.  14.5  km  west  of 
\\'hite  Pass  (north  side  of  SB  12);  46°4()'  X, 
121=31'  W;  elevation  853  m  asl;  April  2000  (8 
specimens  collected  bv  K.  Jcirgensen  and  W.  Leo- 
nard); RBCM  001-00286-001 ;  12  April  2002  (2 
specimens  collected  In W.  Leonard "i;  RBCM  001- 
286-001. 

9.  King  Co..  Mt.  Baker-Snocjualmie  Xational  Forest, 
se\eral  locations  from  0.8  to  1.6  km  west  of  Ste- 
vens Pass;  47°54'  X,  121°06'  ^\";  elevation  1100  m 
asl;  April  and  Mav  2001  (3  specimens  collected 
bv  W.  Leonard  and  W".  XulD:  RBCM  001-00287- 
001. 

10.  Kittitas  Co.,  Wenatchee  Xational  Forest,  Ski 
View;  47°23'  X,  121°22'  \V:  ele\ation  1370  m  asl: 
17  October  2000  (collected  b\  J.  Bauijhi:  RBC:M 
001-00359-001. 

11.  Kittitas  Co.,  Wenatchee  Xational  Forest,  Granite 
Creek:  47°09'  X.  121°05'  W;  elevation  927  m  ash 
19  October  2000  (collected  by  J.  Baugh)  (dissect- 
ed specimen  lost) 

12.  Kittitas  Co.,  Wenatchee  .National  Forest.  Cold 
Creek;  47°20'  X,  121=22'  W;  elevation  80S  m  asl; 
1  Xo\ ember  2000  (collected  b\  J.  Baugh V  RBCM 
001-00361-001. 

13.  Kittitas  Co..  Wenatchee  Xational  Forest.  Mos- 
quito Creek;  47°18'  X,  121°20'  W;  elevation  775 
m  asl:  26  October  2000  (collected  b\-  J.  Baugh); 
RBCM  001-00360-001. 

14.  Kittitas  Co..  Wenatchee  Xational  Forest,  Swamp 
Lake,  47°19'  X.  121=18'  W.  elevation  744  m  as!: 
9  Jtiiv  2001  (collected  h\  J.  Baugh):  RBCM  001- 
00289-001. 

15.  Kittitas  Co.,  Wenatchee  Xati(mal  Forest.  Roaring 
Creek:  47=20'  X.  121=22'  W;  ele\ation  810  m  asj; 
I  1  JuK  2001  (2  specimens;  collected  ii\  J.  Baugh); 
RBC-M  001-00288-001. 


THE  NAUTILUS  116  3  ;95-104.  2(>ii£ 


Pa^f  95 


Xe\\-  species  of  deep-water  Cancellariidae  (Gastropoda)  from 
the  southwestern  Pacific 


Philippe  Bouchet 

Museum  National  d'Histoire  Naturelle 

55  rue  Buftoii. 

75(h:»5  Piuis 

FRANCE 

pbouchet@  cimrs  1 .  mnhn .  fr 


Richard  Petit 

P.O.  Box  3(1 

North  MxTtle  Beach.  SC  29597  US.\ 

r.e.petit@att.net 


.\B.sTR.ACT 

One  new  genus  and  nine  new  species  of  Cancellariidae  are 
described  from  New  Caledonia  from  deptlis  behxeen  2l)()  and 
6(X)  meters.  Thev  are:  Africotriton  mhJphum  new  species.  Mir- 
auilaplwra  new  genus.  MiraudiipJuni  cai/rci  new  species.  Mir- 
audapJwra  maestratii  new  species,  Mcrica  mtiriscti  new  spe- 
cies. Sii-ltia  rocroii  new  species.  Sifhia  spleiulidiila  new  spe- 
cies. SippoiuipJicra  pardalis  new  species.  Siiipoiiaphera  ctj- 
phoma  new  species,  and  Sipponaplwra  goniata  new  species. 
Africotriton  adclphum  new  species  is  the  first  species  in  that 
genus  known  from  outside  South  .\frica  and  .\ustralia.  The  new 
genus  yiirandaphera  is  characterized  b\  its  broad,  non-unibil- 
icate  shell  witli  \er\  lar^e  crenulated  axiiil  ribs,  and  axial  col- 
umella. The  genus  is  composed  of  die  new  species  described 
herein,  Mirandaplwra  macstriitii  new  species  ;uid  \f.  caijrei 
new  species.  ;uid  two  odier  species:  A/,  iosaensis  iHabe.  1961 
new  combination  and  M.  arafurcusi.s  Aerhecken.  1997'  new 
combination,  from  deep  water  off  Japan  and  the  .\rafura  Sea 
respecti\ely.  Triaoiiaphcra  tcramachii  Habe.  1961  ;uid  Aiiatrix 
nodosiharicosa  Petuch,  1979  are  tnuisferred  to  Xippoiiiiphera. 
New  species  of  Merica.  Svrltia.  and  Sippoiuiplicni  are  the 
deepest  dweUing  knowii  representatix es  in  their  respecti\e 
ijenera. 


INTRODUCTIOX 

Tiie  tropica]  deep-water  taima  oi  the  soutliwestem  Pa- 
cific w;is  xiiinuilK  unknown  just  two  decades  ago.  but 
considerable  adxiuices  ha\e  since  been  made  both  in 
tenns  of  sampling  in  die  field  and  ta\ononiic;il  descrip- 
tions ot  the  materiiil  collected.  Empiuisis  h;is  been 
placed  on  die  exploration  ot  the  iu"ea  around  New  Ca- 
ledonia, resulting  in  the  descripdon  of  se\  end  hundred 
new  niolluscan  species  isee.  among  others.  Crosnier  .uid 
Bouchet.  1991:  Bouchet.  1995:  Bouchet  and  .Marshall. 
200U,  .\lthough  the  inventon  is  still  tW  from  c-ompiete. 
it  h;is  ;ilread\  been  revealed  to  be  one  oi  the  most — it 
not  the  most — dixerse  an\-wliere  in  the  world,  with  an 
exceptional  dixei-sification  in  such  families  its  .Seijiienzii- 
dae  (55  species:  .\I;irsIuill.  199U.  Muricitlae  ^90-201) 
species:  Honart.  2001 K  or  Scaphopoila  [73  .species:  SciU"- 
abino.  1995). 


ba.scti  on  tile  literature,  the  tamiU  t^aiici-iianKlac  ap- 
pears poorly  represented  in  the  tropical  South  Pacific-, 
xvitli  h;df  a  dozen  nominal  species  rec-orded  from  shalloxv 
xxater  in  Nexv  Caledoni;i.  to  just  h\o  reported  from  Fiji, 
and  a  single  species  from  the  Socieh  Islands.  Bx  con- 
trast, die  material  originating  from  tlie  recent  expedi- 
tions in  Xexx  C-aledonia.  X'amiatn.  Fiji.  Tonga,  and  \\'allis 
iind  Futiina  contains  about  50  species  of  Cancellariiilae. 
about  -W  ot  xxhich  are  undescribed.  mainlx-  from  depths 
betxxeen  2CX)  and  6tX)  meters.  The  purpose  of  the  pre- 
sent paper  is  to  proxide  descriptions  of  die  more  spec-- 
taciJiir  ot  the  nex\  species  -X  Tiiore  c-omprehensix  e 
monosrapli.  describing  an  ing  the  entire  can- 

ceUariid  fauna  ot  the  regio;..  ox\.  We  restrict  our- 

selxes  in  die  present  paper  to  the  description  of  shell 
moqihologx.  ;md  deter  to  the  sect)nd  article  descriptions 
of  radulae  ixxlien  axailable'  and  discussion  ot  patterns  of 
batlixnnetrical  ;md  geographical  distributions. 

M.\TERLALS  AND  TEXT  CONN  EXTIOXS 

The  nexx-  species  orisjinate  troni  several  dredginsj  pro- 
grams conducted  under  Dn  Bertrand  Richer  cie  F"orges. 
of  Institut  de  Recherche  pour  le  Dex  eloppement  IRD. 
foniierix-  ORSTOMl  Xoumea.  The  cxnilext  oi  the  pro- 
grams. Uiirratixes  of  the  expeditioiis  and  lull  station  lists 
are  presented  bx  Richer  de  Forces  1990'  MISOR- 
STOM  4'.  Richer  de  Forges  1991  iProgranune  L\- 
GOX'.  Richer  de  Forges  1993  S.MIB  5'.  and  Richer 
de  Forges  imd  Chexilion  1996  S.MIB  S.  B.\THUS  2. 
3.  and4\ 

In  the  lists  of  t\pe  and  other  materiid  examined,  in- 
dixidnal  lots  in  NIXTIX  are  unambiguouslx  designated 
bx-  die  combination  of  cruise  acronxni  and  station  num- 
ber. D\\"  refers  to  dredge  hauls.  CP  to  lieam  traxvls:  l\ 
refers  to  live-taken  specimens,  dd  to  emph  shells;  spnis 
to  commerciallx  ontained  specimens,  state  xx-hen 
dredged  not  kniowii. 


specir 


SYSTEM  ATICS 

Fiuiiilx  Canc-ellariidae  Forbes  and  Hanlex.  1S51 
Snbfaniilx-  Plesiotritouinae  Ben  iuid  Max-xvell.  19S7 
C.enus  AfiicotritoJi  Ben  and  .Maxxvell.  I9S7 


Pa-'e  96 


THE  NAUTILUS.  \ol.  lift.  NO.  3 


Figures  1-S.  Xcw  deep-water  C'anccllariidar  l-.'5.  .XfiicdliUdii  inliliiliiiiii  mw  s|H'eics  \.  Il(ilcil\|)c  lici^lil  22,4  iiiiii  scmtli  of 
New  Cialedonia.  SW  of  He  des  Pins.  22°.1T'  S.  I(i7  22'  K.  .590  m.  2.  I'kiIikiiikIi  of  siiccimen  Iroiii  SW  ol  lie  des  I'ms.  22  4.S'  S. 
Ifi7°lf>'  E.  444  -t4.5  in  |HATIIl)S  2:  sla.  D\\7I9|.  3.  Teleoeoiuli  iiiuiiisciilpluic,  s.inie  specimen  a.s  Figure  2,  4-tt.  Miraiidaplwra 

timcstnilii  new  specie.s.  4.  Holotvpe.  Iieiijlit   14.0  ii nlT  nordieastern  eoa.st  of  New  ( .'aletloriia.  Passe  de  llientjliene.  2()".'3;3'  S. 

I64'.5T'  v..  .5:«-61()  ni.  5.  ProtDconcli  of  speenneii  Inmi  Wallis  and  Kntnna.  I.ri9'  S.  17(i°l7'  W.  .3.50  ni  [Ml!S(  )R.ST()M  7:  sta 
D\\fi01|.  6.  Teleoeoneli  niierosenlptnix-.  same  specimen  as  Fii;ni'e  .">.  7-8.  Minniildphcni  ciii/iri  new  species.  7.  Specimen.  Iniiilil 
27.3  nnn:  norlli  of  .\ew  Caledonia.  UfOT  S.  I(i3°13'  K.  .34I-.3.5I  ni  iB.VriirS  4:  sla.  in\»021,  S.  Ilolotvpe.  Iieiijht  .3S.7  mm; 
South  of  New  Cal.-donia.  off  Passe  de  Kouare.  22" W  S.  Ui6  l.'j'  K.  .3(lll-.)70  m.  Se.ile  Imes  -    100  |i.m  (3.  (Si  and  .'lOO  |jlui  (2,  .5). 


Type  Specie.s:     Epiilnmiiis  (■rchrilinitiis  i,.  15. 
Sowerbv  III,  190.3.  In  orit^inal  designation. 

Africotrilon  adclpiiiitn  new  species 
(Figures  1-3' 

T\pe  Material:  Ilolotvpe  (22.4  mhii  lieiglit  X  7.S  iiiiii 
width)  (1\)  (Figure  I)  and  3  paratopes  (ddi.  all  from  the 
t)pe  localitA.  in  MNHN. 


'INpe  Loealilv:  .South  ol  Nt'w  (^aK'douia.  SW  ol  lie 
di's  Pins.  22°47'  S,  lfi7°22'  P].  .390  ni  |  MUS(  )HSr(  )\l 
4:sla.  nW22fil, 

Oilier  Material  Fxainined:  South  ol  New  Caledonia. 
HlOCAl.:  sta  1)\\44.  22  47'  S.  Ifi7"l4'  E.  440-4.50  ni, 
2  ju\.  K.  I  in\.  (I(L-  S.MIH  2:  sla.  DW'l,  22°53'  S. 
167'  I. r  E.  43S-444  in.   I   jii\.  K.— S.MIB  3:  .sta.  D\V2fr 


p.  Bnuclirt  and  R.  Petit.  2002 


Pasie  97 


22"55'  S.  167"16'  E,  450  m.  1  K.^BATlllS  2:  sta. 
DWTUJ.  22°4S'  S.  lfi7°lH'  E.  444-443  m.  4  K  iFi^uivs 
2,  3). 

Description:  Protde-oiu'li  iiiaimiiilatf.  paiii'ispiial.  of 
1,1  whorls,  diameter  1.25-1.30  mm  (Figure  2).  Traii.si- 
ticiii  to  teleoconeh  marked  h\'  oii.set  of  closeK  .spaced 
spiral  cords.  Teleoconeh  ol'  sexen  moderatcK  ronnded 
whorls;  spire  angle  2fi'-27^  Prominent  ortliocline 
rounded  \arices  tliat  extend  al)o\e  the  appressed  suturi' 
are  formed  irregularK.  hut  nsualK  at  ahout  240"  inter- 
vals. Spiral  sculpture  ol  over  (SO  low  closeK  spaced  spiral 
cords.  ."Vxial  sculpture  of  widel\  spaced,  rounded  ncm- 
collaiiral  rihs,  7  between  final  two  varices,  witli  closeK' 
packeil  microscopic  growth  lines  (Figure  3).  .\perture 
narrowK'  o\"ate.  Outer  lip  with  weak  crenulations  on 
edge  hut  smooth  within.  .\  shielddike  callus  produced 
on  the  inner  lip.  weak  ovi-r  the  parietal  area  h\it  well 
produced  oxer  the  imluctural  area  and  extending  o\'er 
the  siphonal  constriction.  Columella  ahiiost  axial  with  a 
narrow  fold  at  the  center.  A  second.  \en  weak  fold  pos- 
terior to  the  center  fold  is  xisihie  on  some  specimens. 
Siphon. il  canal  narrow,  opt'ii,  strongK  recnnetl.  Shell 
color  cream  to  vellow-hrown  with  fi  or  7  red-hrowii  spi- 
ral bands,  narrow  on  small  shells  but  wiile  on  some 
adults,  appearing  as  weak  hands  except  on  the  tops  of 
the  varices  where  the\-  appear  as  narrow  stripes. 

Etvmolog)-:  The  Greek  ddcliihos.  brother,  utilized  to 
highlight  the  resemblance  of  the  species  to  others  in  the 
genus;  treated  here  as  a  Fatin  adjective. 

Remarks:  Of  the  si.\  species  inchuled  in  this  genus  when 
introtluced,  five  are  from  South  Africa  and  the  sixth,  .A. 
carifiiipcx  Beu  and  Max-xvell.  19S7,  is  from  off  New 
South  Wales,  Australia.  The  new  species  differs  from  A. 
carinapex  in  being  much  more  slender  with  less  convex 
whorls,  and  in  having  stronger  spiral  cords  and  weaker 
axial  ribs.  Africotriton  mlelphum  resembles  more  the 
South  African  A.  Mlhurni  Beu  and  Maxxvell,  1987.  but 
that  species  has  more  prominent  spiral  cords  and  weaker 
axial  ribs. 

Subfamilv  Cancellariinae  Forbes  and  Hanlev.  1S51 
Genus  Mirandaphcra  new  genus 

Type  Species:     MiraiuUiphcrd  c<np-ii  new  species 

Description:  Shell  rehitivelv  large,  reaching  over  35 
mm.  Teleoconeh  of  7-9  whorls.  Shell  somewhat  atten- 
uated vvitli  large  axial  ribs  that  dominate  the  shell  sculp- 
ture. Ribs  angk'd  or  loimdcd  at  the  shoulder  over  wliicli 
tlun  project  ;is  sm;ill  coi'onations  but  rarelv  extend  hack 
to  the  sutmc.  Sutinal  ramp  depressed.  The  sutiuc  is 
onlv  slightlv  impressed.  Small  nodes  are  formed  where 
weak  spiral  cords  cross  ribs.  Spiral  cords  are  nnich  stron- 
ger anil  more  evitlent  on  the  ribs  than  in  the  interspaces. 
Outer  lip  tliick.  slightlv  flared  on  outer  edge  and  ex- 
tending back  over  tlie  paiietal  wall  as  a  rounded  shield 
that  extends  out  at  tlie  imluctural  area.  No  stromboid 
notch  visible   on  outer  lip.   Columella  axial  with   two 


strong  folds  and  a  thiid  fold  which  slls  on  the  ediie  of 
the  siphonal  fold  before  turning  avvav  to  parallel  other 
folds.  Folds  do  not  extend  out  to  the  edge  of  the  iiidue- 

Inral  shelf   I  nibilicns  absenl. 

Discussion:  ik'sides  the  twd  new  species  described 
here,  we  include  Tiitidiuiphcrii  lo.sdciisis  Ilabe,  1961  {re- 
centlv  |il;iced  in  Sohilia  In  llasegavva,  2001:  5S3)  and 
Solatia  arafiirciisis  N'erheckeu.  1997  in  Mirandaphcra 
The  resemblance  of  these  t;ixa  to  Si>latia  is  onlv  ven 
superficial.  The  tvpe  species  of  Solatia.  S.  piscatoria 
(Gmelin.  1791  ).  has  weak  colimiellar  folds,  an  apjiressed 
suture,  and  a  flaring  apertmc  among  otiier  differences. 
Species  ol  Miunidaplwra  iiave  a  superficial  resemblance 
only  to  high-spired  species  ni'  Scalptia  Jousseaume.  1SS7 
from  which  diev  ditler  in  having  an  elongated  tabulate 
shell  with  vvi-v  large  crenulated  axial  ribs  and  a  axial  col- 
umella Minindaphcra.  as  understood  here,  is  a  deep- 
water  genus.  No  fossil  spi-cies  luivc  been  reco^jni/ed  in 
the  literature. 

Et\'moloii;\ :  The  Fatin  iiiiraiidus.  nn'aning  wonderful 
Ol'  sli'iinge.  eoinbined  with  aplwra.  Apliera  was  intro- 
duced as  a  g(.'uus  in  Cancellaiiidae  and  later  used  as  a 
stem  name  for  various  cancellariid  genera,  all  treated  as 
feminine.  A  search  for  its  origin  reveals  onlv  a  few  us- 
ages, the  earliest  of  which  are  as  the  given  name  of  wom- 
en in  Eni'Iand  in  the  mid-17'''  C'entnn. 


Mirandaphcra  cai/rci  new  species 
(Figures  7-S) 

T%pe  Materia!:  Ilolotvpe  i3S.7  mm  height  X19.3  nun 
width)  ilv  1  in  MNllN  i  Figure  Si. 

Type  Locality':  South  of  New  Cafedonia.  oft  Passe  de 
Kouare.  22"49'  S,  166°45'  E,  300-370  m  [B;VTHUS  2; 
,sta.  D\\'7311. 

Other  Material  Examined:     North  of  New  Caledonia. 

B.-VTllUS  4;  sta.  1)W9()2,  19°()1'  S.  fa3=15'  E.  341-351 
m.  2  dd  (Figure  7i. — l-'rom  conunercial  sources,  boat 
Tui  IF  off  I^elep  Islands.  3  spms. 

Description:  Protoconch  smooth,  of  1.3  whorls,  diam- 
eter S30  [jLui.  Transition  to  teleoconeh  marked  bv  strong 
axial  ril).  Teleoconeh  of  about  9  elongate  whorls.  .■\.xi;il 
sculpture  of  strong  rounded  ribs,  about  12  on  bodv 
whorl  and  14  ou  penultimate  whorl.  The  ribs  are  bian- 
gular  on  the  shoulder  over  which  thev  recurve  and  ex- 
tend partialK  onto  the  othenvise  smooth,  fiat  sutural 
i'am|).  Sulni'e  barelv  impressed.  S]')iral  sculpture  of  about 
S  priman  eoi'ds  that  form  prominent  noiles  wiiere  they 
cross  the  iixiai  ribs.  Nodes  strongest  on  (he  shoulder. 
wide,  biangular  .\bout  4  priman  spiral  cords  visible  on 
penultinuite  whorl.  Numerous  fine  secondarx  spiral 
cords  cover  the  shell.  S  or  more  between  each  pair  of 
priinan  cords  with  an  occasional  si'condarx  spiral  cord 
intermediate  in  strength,  not  intersected  by  axial  riblets. 
Ribs  on  adult  shells  sometimes  formed  as  thickened  var- 
ices with  subse(juent  growth  originating  from  under  the 


Page  98 


THE  NAUTILUS.  W 


IH,  No.  3 


outer  lip.  Aperture  elougate.  Outer  lip  tliick,  .siiiootli. 
with  a  narrow,  smooth  shelf  inside  the  lip.  About  12 
strong  lirae  extend  from  lip  onlv  a  short  way  into  the 
aperture.  A  posterior  canal  is  formed  under  the  shoulder 
and  is  further  delineated  In'  a  pustule  on  the  parietal 
Willi.  Shield-like  parietal  callus  well-developed.  Colu- 
mella with  three  strong  folds,  the  anterior  one  weakest 
and  bordering  the  distinct  siphonal  fold  before  turning 
to  parallel  the  other  folds.  Folds  do  not  e.xteud  to  the 
end  of  the  inductural  Ciillus.  Body  whorl  onlv  slightK 
constricted  behind  the  siphonal  fasciole.  Umbilicus  ab- 
sent. Shell  color  off-white  or  beige:  prinian'  spiral  cords 
slightK'  darker,  light  \ellow-bro\\Ti. 

EtjTnolog>:  Named  for  Dr.  Patrice  Ca\Te,  head  of 
IRD's  Department  des  Ressources  \'i\  antes,  in  recog- 
nition for  his  support  to  ta\onom\'  and  biodiversity'  ex- 
ploration. 

Remarks:  Minnulaphcra  cai/rci  differs  from  M.  to- 
saeiisis  (Habe,  1961)  and  M.  (mifiireims  (\'erhecken. 
1997)  in  having  strong  lirae  within  the  aperture  and  a 
more  turreted,  attenuate  shell  on  which  the  spiral  sculp- 
ture is  prominent  in  profile.  Miranchiphera  maestratii 
has,  among  other  differences,  more  numerous  axial  ribs, 
stronger  spiral  sculpture,  and  a  more  rounded  profile. 

Miniiulaplwra  iimc'-itratii  new  species 
(Figiu'es  -f-6) 

T\pe  Material:  Holohpe  (14.0  nun  height  X  8.4  mm 
width!  idd'  in  .\1NHN  (Figure  4). 

Type  Locality:  Olf  northeast  coast  of  New  Caledonia, 
Pas.se  de  Ilienghene,  20°33'  S,  164°57'  E,  533-610  m 
[BATH US  4:  sta.  D\V948]. 

Other  \taterial  Examined:  \anuatu:  MUSORSTOM 
S:  sta.  DWlOfil.  Krl5'  S.  i67°20'  E.  458-512  m,  1  dd; 
Fiji:  MUSORSTOM  10:  sta.  CP1341,  16°52.5'  S, 
177°43.7'  E.  500-614  m,  1  dd;  Wallis  and  Futuna:  .MU- 
SOR.STOM  7:  sta.  D\\Ti01,  13°19'  S,  176°17'  W,  350  m, 
1  dd  (Figures  5.  6), 

Description:  Profoeonch  prominent,  of  1.3  whorls,  di- 
ameter 850  [xm  (Figure  5).  Onset  of  teleoconch  markeil 
by  a  strong  axial  rib.  Teleoconch  of  about  6  whorls  or- 
namented with  high  narrow  axial  ribs  barcK  wiiler  than 
interspaces.  The  axial  ribs.  14-16  in  number,  extend  over 
the  rounded  slioukler  and  turn  down  to  a  narrow  sutural 
ramp,  but  do  not  extend  all  the  wax  to  the  moderat(>K 
impres.sed  suture.  Spiral  sculpture  of  narrow  tleu.s(>l\ 
packed  spiral  coi'ds  l^liat  become  denser  and  increase  in 
height  at  regular  intenals,  forming  bead-like  nodes  on 
the  ribs.  There  are  8-10  nodes  on  the  bod\  whorl  ribs 
with  approximately  16  cfirds  bcKveen  centers  of  nodes. 
Spiral  cords  intersected  In  finer  incremental  riblets  that 
form  an  intritacalx-like  microsculpture  (Figure  6).  I'inal 
rib  is  produced  as  a  terminal  \arix.  Aperture  elongate, 
narrow.  Outer  lip  thickened  with  8  lirae  within  that  ex- 
tend to  the  outer  edge  btit  do  not  tiescend  dei^pK  into 


the  aperture.  Posterior  can;il  is  formed  under  the  sh(ml- 
der  and  is  further  delineated  b\-  a  pustule  on  the  parietal 
wall.  Parietal  callus  well-developed  but  not  extending 
out  as  a  shield.  Columella  with  three  strong  folds,  the 
anterior  one  weakest  and  bordering  the  siphonal  told 
before  turning  to  parallel  the  other  folds.  Folds  do  not 
extend  to  the  outer  edge  of  the  inductura.  Bod\'  whorl 
slightlv  constricted  behind  the  small  but  w^ell-defined  si- 
phonal fasciole.  Uml)ilicus  absent.  Shell  color  light 
brow'nisli-\'ell(n\. 

EtjTnoIogj':  Named  lor  Philippe  Maestrati,  .MNHN 
museum  technician,  in  appreciation  for  his  dedication  to 
processing  and  sorting  much  of  the  New  Caledonia  ma- 
terial. 

Remarks:  Miraiulaplwra  maestratii  is  separable  from 
M.  caijivi  h\  its  closeK  spaced  axial  ribs.  The  spiral  cords 
of  M.  maestratii  are  also  unique,  being  so  closelv  spaced 
tliat  the  interspaces  appear  as  incised  lines.  The  delicate 
pattern  the\'  form  is  especialK'  noticeable  when  there  is 
a  \ailx  on  which  the\'  are  \isible. 

Genus  Merica  H.  and  .-K.  Adanis,  1854 

T\'pe  species:  CanccUaria  mclanostoma  Sowerby, 
1849,  b\'  subsequent  designation  of  Cossmann  (1899). 

Merica  maiisca  new  species 
(Figm-es  9-12) 

T\pe  Material:  Holotvpe  (25.7  mm  height  X  18.0 
nnii  width)  iK  )  (Figure  9)  in  MNHN. 

T>'pe  Locality:  ( )tl  northeast  coast  of  New  Caledonia, 
Passe  de  Hienghene,  20°34'  S,  164°58'  E,  470-190  m 
[BATHUS  4  sta.  CP947]. 

Other  Material  Examined:  New  Caledonia:  BA- 
THUS 4:  sta  CP889,  2r01'  S,  164°27'  E,  416-433  m, 
1  dd  (Figure  12).— HALIPRO  1:  sta  CP851,  21°43'  S, 
166°37'  E,  314-364  m,  1  Iv  (Figure  11):  Vanuatu:  MU- 
SORSTOM 8:  sta.  CP1136,  15°41'  S,  167°02'  E,  398- 
400  m,  1  dd  (Figure  10). 

Description:  Protocoucli  smooth.  shin\.  with  1.0 
wliorl  iFiguri'  12).  Transition  to  lcleoconi-li  marked  In' 
a  slight  depression  and  onset  of  axial  and  spiral  sculp- 
tures. Teleoconch  of  up  to  6  rovmded  whorls.  Shoulder 
small,  rounded,  sloping  back  to  an  impressed  suture 
lorniing  a  narrow  channel.  Axial  scul[)ture  of  about  1-1- 
16  low  axial  ribs  on  b()d\  whorl  of  adult;  slightlv  more 
on  earlier  whorls.  Ribs  rounded  in  profile  and  narnnver 
than  intencTiing  spaces.  Periodic  inti-rnal  \arices  (see 
Harasewxch  and  Petit,  1982:  ill),  noticeable  on  outer 
surface  of  shell  as  thickened  axial  ribs,  each  followed  by 
a  relati\('l\'  flat  area.  Spiral  sculpture  oi  strong,  broad 
cords  ( 16-20  on  bod\'  whorl,  6-7  on  pemiltimate  whorl); 
those  on  shoukk-r  cnnvded.  others  usualK  with  a  sec- 
ondan  cord  in  the  interspaces  and  rareK  a  tertian  cord. 
.\pcrture  lai'iif  with  ;i  wideK  elliptical,  prosocline  outtn" 


p.  Bouclu't 


R.  Petit.  2002 


Page  99 


Figures  9-17.      Xe«  deep-water  Caneellariulue.  9-12.  Mcrica  mari.sca  ne^^  spec.es.  9.  1  l..l..t>pe.  l,ei>.|,l  2.yT  mmu,  oti  ,K>rtheasten. 
coast  of  New  Caledonia.  Passe  de  HienaliiMie.  2(f34'  S.  164°5S'  E.  470-490  >n.  10.  Specn„e„  Iron.  Xu.u.atn,  he.gl.t  -  1 ,  .9  .....K 

MUSORSTO.M  S:  sta.  CP11361.  II.  Specin.en  troni  New  Cidctlon.a.  I.eiglit  IS.,  mm:  SI  45   s. 

12.  Protoconcli  of  specimen  fiom  off  the  western  coast  o(  New  Caledonia. 

CP.S891    13-14.  Sieltiii  ivrroii  new  species.  13.  Ilolot\pe.  I.eiglit  14.3  mm: 

Onest.  23°20'  S.  16S°01'  E.  .501-365  m  [B.\TiIUS  3:  sta.  D\\S.30].  14. 


15°41'  S.  167°02'  E.  39S-400  m 

166°3T'  E.  314-.364  m  [HALIPRO  1:  sta.  CPS.51] 

21°01'  S.  164°27'  E.  416-433  m  [B.-\THUS  4:  sta 

south  of  New  C:aledonia.  Xoifolk  Riilge.  Banc  Jumean 

Protoconc 

15-1 

167 

23' 

Sc;ile  line 


lull  ui   .\e\\    \..aie(.UMiia.   -\vhiuii\   iiivii^^r.   i.»txiis.      uiiiv^n.   ^..v....  —  —  .  ri..i/-A/"«i        i       i^M'O" 

otoconch  of  specimen  fron.  sonth  of  New  Caledonia,  S\\-  of  He  des  Pi.is.  23=0()'  S.  16,  16  E.  .5o(  -n  [BIOC.VL:  sta.  D\  3,]. 
1-17.  Svcltia  splculklulu  ,iew  species.  13.  Holot^pe.  height  11.0  mm:  so.,th  ol  New  Cak^lonia.  S\\  of  He  d.-s  I  ins.  oO  S. 
i7^2P  E  500-.504  m  [SMIB  8:  sta.  D\\-2(l]].  16.  Parat\pe.  height  10.1  nm.:  south  ol  New  (.aledcmi:..  S\\  ot  He  des  I  ms.  -  o9  - 
;°00'  S    167°21'-I67°23'  E   491-558  m  [SMIB  8:  sta    i:)\\lH.3-196l.  17.  Protoco.ich:  specime.i  Worn  s:mie  local.t^  as  l-ii;nre  Hr 


.500  M-m  (12.  14.  17). 


Page  100 


THK  XAITIIAS,  \<)1.  llfi.  No 


lip.  Outer  lip  with  a  HiuK  stTiati'd  i'(l<ic;  inner  surface 
witli  14  spiral  lirae  that  (iiniinish  aliout  a  (|uarter  whorl 
into  the  aperture.  C'olnnulla  with  ■>  folds,  the  posterior 
one  being  slightK  larger  and  the  anterior  one  torniiug 
the  edge  of  the  short,  narrow  canal.  The  anterior  lold 
extends  to  the  edge  of  the  inductural  callus  with  smaller 
fold-like  features  or  pustules  sometimes  present  on  out- 
er edge  of  inductnra.  Color  \ellow-hro\\ii  with  a  dark 
red-hrown  hand  at  shoukler  and  smallei'  lines  or  hands 
on  l)od\  whorl.  \  white  hand  is  sometimes  present  just 
anterior  to  the  shouldei'  ami  just  anterior  to  the  periph- 
ery. 

Etvinologv:      Latin  iiiaii^ca.  f .  a  large  t\pe  of  fig. 

Reniark.s:  Mcricd  piniiii  has  many  features  in  com- 
mon with  the  Philippine  Merica  ekti/phos  Petit  and  Har- 
asew\ch,  2(100.  which  has  stronger  sculpture  with  lieavy 
nodes  formed  at  intersections  of  spiral  cords  and  ;L\ial 
rilis.  In  addition,  M.  ckfi/plnis  has  a  roundetl  and  more 
solid  shell,  and  the  protoconch  is  '4  whorl  larger  than  in 
M.  piniin.  Species  of  Merica  li\e  suhtidalK'  and  on  the 
continental  shelf  The  batlnal  M.  piniiii  is  apparentK  (he 
deepest  living  species  in  the  genus. 

Cienns  Svcltiii  [oussi'anme.  1SS7 

Type  Species:  "Svclliti  i(nico\<i  [sic|  15rocc."  [=  Vol- 
uld  rarricosa  Broci-hi,  fSf4l,  h\  original  di'signatiou. 

Svcltid  rocroii  new  species 
(Figures  1:5-14) 

T\pt'  Material:  Holot\pe  (14.3  mm  height  X  S.9  nun 
width !  iK  1  (Figure  13)  and  one  paratope  (dd)  from  the 
t\pc  locality  in  MNHN. 

Type  Locality-:  South  of  New  Caledonia,  Norfolk 
Ridge,  Banc  jumean  Ouest..  23°20'  S,  lfiS°Of'  F,  3fif- 
365  m  |B.\TIIUS  3:  sta.  DWS30|. 

Other  Material  Examined:  New  C;aledonia:  BiO- 
CAL;  sta.  D\V.37,  2.3°()()'  S.  IfiT'lfi'  E.  350  m.  1  ju\.  K. 
(Figure  14),  1  jnv.  dd.— MUSOKSTOM  4;  sta.  DW'lSl. 
18°57'  S,  163°22'  E,  350  m,  1  dd.— BATHUS  4:  sta. 
D\V931,  1S°55'  S,  163°24'  F,  360-377  m,  1  dd.— From 
connnercial  sources,  boat  Tui  II,  off  Belep  Islands,  3 
spnis. 

Description:  Protoconch  smooth,  erect  of  I  O  whorl. 
diameter  !  ..30  mm.  axis  of  protoconch  tilted  to  teleo- 
conch  (Figure  14(.  Transition  to  teleoconcli  eleark 
marked  by  a  strong  \arix-like  line,  angled  at  the  shoulder 
and  followed  b\-  onset  of  axial  ribs.  Teleoconcli  of  about 
five  whorls,  .\\ial  sculpture  of  low,  wideK  spaced  libs, 
se\en  on  body  whorl  and  ten  on  |iennltiiriate  whoil.  ex- 
tending from  the  suture  to  the  ba.se  ol  the  shell  Spiral 
sculpture  ol  one  cord  situated  niidwax  on  whorl  and 
forming  an  angled  shoulder  w  ith  sharp  short  sjiines  |m'- 
sent  at  intersections  with  iLxial  ribs.  A  weaker  spiral  cord 
is   .sometimes   present   betvxeen   the   shoulder   and   the 


slightK  impressed  suttu'e.  About  fi\c  weak  spiial  colds 
are  located  anterior  to  the  shoulder,  the  first  anterior  one 
widely  spaced  from  shoukler  and  forming  a  slight  angle 
to  the  bod\  whorl  Remaining  spiral  cords  weaker  and 
more  clost'K'  spaceil.  ,\pertnri'  large,  o\al.  Outer  lip  pro- 
socline.  formed  b\'  a  terminal  \arix,  smooth  within.  Pa- 
rietal wall  with  weak  callus,  becoming  stronger  and 
slightK'  reflected  at  the  inductnra.  Columella  with  three 
strong  folds,  the  anterior  one  forming  the  edge  of  the 
short  siphonal  canal.  Shell  thin,  white,  translucent. 

Et\iiiolog\':  Named  for  M.  )ean-Paul  Rocroi.  .\INHN, 
in  appreciation  ol  liis  contiibutiou  to  inollnscan  iioinen- 
clature. 

Remarks:  The  new  species  ina\  be  cmnparetl  to  oiiK 
one  Recent  species  other  than  Svcltia  splendidula .  de- 
scribed herein,  is  S.  'gladiator  (Petit,  1976),  which  occurs 
off  the  Cialapagos  Islanils  at  a  depth  of  200  m.  Although 
their  protoconchs  are  entirelx  different  (the  protoconch 
ol  S'.  f^Iddiator  is  nmltispiral  indicating  planktotrophic 
lar\al  dexelopment),  the  two  species  share  the  "window- 
like" sculpture  below  tlie  shoulder  formed  h\  the  wideK' 
spaced  cords  and  ribs.  The  spines  on  the  shoulder  of  .S. 
nx-niil  are  much  shorter  than  in  S.  fjjadiator.  The  Elu- 
ropean  Tertian  species  Calcarata  calcarata  (Brocchi. 
1S14)  is  startliugK-  like  S.  rocroii  in  appearance  but  that 
species  differs,  among  other  features,  in  being  slightK' 
umbilicate. 

Sieltia  splciididiild  new  species 
(Figures  15-17) 

Type  Material:  llolot\pt'  ill.O  mm  height  X  6.9  mm 
widtli)  (K)  (Figure  15)  and  three  paratxpes  (1  l\.  2  dd( 
in  MNHN. 

Type  Locality:  South  of  New  (Caledonia.  SW  of  Ik' 
d(''s  Pins,  22°5b'  S,  167°21'  E,  500-504  m  [S.MIB  S:  sta. 
l)\\'20fj. 

Other  Material  Examined:     South  of  New  Caledonia. 
SW  of  He  des  Pins;  SMIB  S:  sta.  D\\I93-196,  22°59'- 
23°()0'  S,  167°2r-167°23'  E,  491-558  m,  1  Iv,  2  dd 
iparatspes:  Figures  16.  17). 

Description:  Protoconch  smooth,  erett.  ol  0.9  whorls, 
dianu'tei'  1.20  mm,  axis  of  j^rotoconcli  tilted  to  teleo- 
conch  (Figure  17'.  Transition  to  teleoioiu-h  inaikeil  li\ 
the  appearance  ol  an  axial  rib  with  angled  shouldei'.  Te- 
leoconcli of  .3+  lapidK  expandinti  whorls.  Axial  sciilp- 
(iirc  ol  eveiiK  spaced  sharp  ribs,  about  12  on  jioiK  whoil 
that  ai'<'  contiiinoiis  Ircnii  the  suture  to  the  base.  I'he 
libs  are  produced  sli'.^lilK  above  and  over  siibseijneiit 
grow  til.  .Spiral  sciilplme  ol  about  seven  sliaq)  sjiiral 
cords.  (  )ne  cold,  soiiiew  ha(  stronger.  I  onus  the  shoulder 
over  w  I  lid  I  the  a\ial  ribs  cross  ,md  angle  back  to  a  sliglit- 
K  impressed  suture.  .\  second  strong  coril  sometimes 
|)reseiit  at  line  of  sntiiral  attachment.  On  the  final  por- 
tion ol  the  bodv  whoil  the  ainj;led  shoulder  becomes  ob- 
solete and  the  ribs  eiiivc  siiiootlilv  b.ick  to  an  impressed 


F.  I^oiiclict  and  R.  Petit.  2002 


Page  101 


Fissures   18-27.     New   ilccp-watiT  ( laiici'lluriidac.   18-21.  Mjtjxniiiiilirin  pariliilis 
nuitli  ol  New  C:aleaiinia.  CrMul  l'assa-;r.  19  11(V  S,  16.y29'  K.  230  m  [MrS()HST( 

mm:  north  of  New  Caledonia.  Craiul  Passaiic   19  Di     >.  loo  .>ii    I'..  — ^i  m  |  i.av..v_;.\.  Ma,   ii-^ji^-  '  ,  ..  i  i 

-.rid  111  ISMIB  S;  sla.  1)\\  ISl].  21.  .Mieroseiilptnrc  ol  teleoeoncli; 

)rtli  ol'  New  CaledonI; 


/(A  m-w  >|H-eK'S.  18.  l-araUpr.  hciul.t  20.0  iiinu 
lOM  4:  sta.  D\\1S5|.  19.  liolotvpe.  lieii;lil  2.5.5 
ON    sta    114S1.  20.  rrotoeoiuii:  spceiiiien  IVoiii 


South  ol  New 


Scale  lint 


10(1  fjLiii  !21.  24,  27'  ami  .500  ijliii 


Pa2c>  102 


THE  NAUTILUS.  Nol.  1  Ifi,  NO.  3 


suture.  Thf  .spiral  cords  do  not  cross  over  the  a.\ial  ribs. 
Aperture  large,  ovate.  Outer  lip  prosocliiie,  tliin,  without 
internal  lirations.  Parietal  area  without  callus.  A  small 
callus  is  sometimes  formed  at  \er\  base  of  intluctura. 
Columella  with  three  folds,  the  posterior  one  larger  and 
descending.  The  two  anterior  folds  situated  at  ends  of  a 
shelf-like  structure  bordering  the  short  siphonal  canal. 
Bodv  w'horl  sliglitK-  constricted  behind  the  weak  siphon- 
al fasciole.  .Shell  color  white,  translucent. 

Et\Tnolog\':  Dimimitixc  of  the  Latin  adjectixc  sjilcii- 
(lidiis  [-U.  -HuiK  meaning  show^■  or  magnificent. 

Remarks:  Sicltia  s])hn(lid\d(i  closeh'  resembles  S'.  ro- 
croii  from  which  it  differs  in  possessing  more  numerous 
and  stronger  spiral  cords  and  a\ial  ribs,  and  in  ha\ing  a 
much  less  angular  shoulder.  The  unusual  combination  of 
sculpture  and  translucence  gi\es  the  shell  a  shoji-like 
appearance.  The  bathvmetric  distribution  of  the  geuus 
Sveltia  is  unclear  due  to  the  confused  generic  allocation 
of  included  species,  but  Sveltia  splcndkluhi  is  apparentK- 
the  deepest  dwelling  species  in  the  genus. 

Genus  \ij)po)iiii)licra  Ilabe,  1961 

T^pe  .Species:  Xippiiii(ii)licni  Itnhci  Petit,  1972,  In 
iC.ZX  Opinion  1052, 

Nipponaphcrn  fxinldlis  new  species 
(Figures  1S-21> 

Type  Material:  Ilolotvpe  (25.5  m  height  X  IS.O  nun 
width)  (Iv)  (Figure  19)  and  one  parat\pe  (Iv)  (Figure  18) 
in  MNHN. 

Type  Locality:  North  of  New  Caledonia,  Grand  Pas- 
sage, 19'()7'  S,  ](i:r:W  K,  220  m  [LAGON:  sta.  114S]. 

Other  Material  E.xaininecl:  New  Caledonia:  .\1U- 
SORSTOM  4:  sta.  D\\T85,  19°0(i'  S,  lfi.3°29'  E,  230  m, 
I  1\-  (parat\pe.  Figure  IS).— Sta.  DW227.  22°46'  S. 
167°20'  E,  300  m,  1  (kl— SMIB  5:  ,sta.  DWll.  23°42' 
S,  168°0r  E,  400  m.  1  dd  [worn;  allocation  uncertain].— 
SMIB  8:  sta.  nW'ISl,  23ns'  S,  lfiS''05'  E.  311-330  m, 
1  juv.  Iv  (Figures  20,  21).— Sta.  1)\\  1S2-1S4,  23°18'- 
23°19'  S,  168°05'  E,  305-367  m,  1  (kl.^BATHUS  2: 
sta.  D\V714,  22°38'  S,  I67°10'  !■:.  121  m,  1  (kl,— BA- 
THUS  4:  sta.  D\V942,  19°04'  S,  163°27'  E,  264-270  ni, 
1  juv,  1\. — From  commercial  sources,  boat  Tui  11,  off 
Bele|)  Islands,  3  spms. 

Description:  I'rotoconcii  corrodid  but  appaicniK  ccm- 
.si,stiiig  of  about  1.5  whorls,  teleoconch  of  5.2  whorls. 
Spire  high.  Sculpture  of  imbricated  axial  lam(>llae  and 
raised  spiral  cords  that  cross  to  form  S(|narish  intenals. 
Aperiodical  axial  \arices,  six  on  final  whorl.  fi\c  on  pen- 
ultimate whorl,  and  two  on  antepenultimate  whorl.  Spi- 
ral sculpture  of  about  10  spiral  cords  on  peimltimale 
whorl  and  about  20  on  final  whori  Second  oriler  sculp- 
ture of  iTicriMuental  lines  crossing  spiial  threads  and 
forming  fiucK  reticulate  sculptiuc  in  the  s(|narish  inter- 


vals (Figure  21 ).  Spire  whorls  regularly  convex,  gradualK' 
l)ecoming  angular  at  shoulder  on  final  two  whorls.  Su- 
ture deepK'  impressed.  Both'  whorl  onl\  slightK  con- 
stricted behind  siphonal  fasciole.  Outer  lip  thin,  smooth 
innnediateh  within  but  then  with  about  18  lirations  that 
extend  deepK  into  the  aperture.  Inner  lip  reflected  as  a 
parietal  shield  partly  extending  over  the  umbilical  area 
in  the  inductural  region.  Columella  slightlv  concave  witli 
three  folds,  the  anterior  two  folds  close  together,  unich 
like  one  large  bifiu'cate  fold.  Deep  umbilicus  well  de- 
veloped. Siphonal  canal  short  but  distinct.  Shell  liack- 
ground  color  cream\-wliite  with  brownish  blotches  on 
the  periphen'  of  the  whorls  behind  \arices;  priman'  spi- 
ral cords  brown  on  the  varices. 

EtymologA':  The  Latin  panlalis.  a  female  panther.  To 
be  treated  as  a  noun  in  apposition.  Selected  because  of 
tile  brown  blotches  behind  the  varices. 

Remarks:  The  protocouch  of  the  holot\pe  is  corrod- 
ed. The  protoconcli  of  the  paratxpe  consists  of  1.8 
whorls,  diameter  1.15  nun  (Figure  20).  Among  the  spe- 
cies of  Nipponapltcra  known  at  present,  this  new  species 
most  resembles  .Y.  tciriiiuicliii  (Habe,  1961),  new  com- 
bination, which  is  more  distinctly  angled  at  the  shoul- 
(lei;  and  has  coarser  sculpture,  (The  transfer  of  Tr/go/i- 
iiplicra  fcriiiiwchii  to  Xipjioiiaphcra  will  be  discussed  in 
oui'  later  monograph. ) 

Nipponaplwni  ci/phoniti  new  species 
(Figures  22-24)' 

Type  Malei-ial:  ilolotxpe  (13.1  height  X  9.2  nun 
width)  (ddl  (Figures  22-24)  in  MNHN;  2  panit\pes. 
Petit  collectif)n. 

Type  Locality:  North  ul  New  C^aledonia.  (irand  Pas- 
sage, 18°55'  S'  163°24'  E,  350-365  m  [HALICAL  1;  sta. 
n\\04]. 

Other  Material  E.vaniinecl:  From  rommei'(i;il  sourc- 
es, off  Hel(-p  Island,  dredged  :it  400  ni  In  bo;it  Tn  II, 
M;l\   2001,  2  sjims  (p;u";lt\]n's). 

Description:  Protocouch  glassw  white,  with  0.9 
whorls.  dl;nneter  0.93  mm  (Eigiu'e  23).  Transition  to  te- 
leoconch marked  In  ;i  small  \;nix-like  axial  rib  followed 
b\'  spinil  ;md  ;ixi;il  scnlptuic.  Spiral  sculpture  of  flat  cords 
ol  xiUAiiig  sizes,  i'rimarv  cords  ni)ini;ilK  separated  b\' 
slightlv  smaller  seconilan  cortls  with  even  smaller  ter- 
tian cords  in  iuters|i;ices  between  the  jiriman-  and  see- 
ondiUA  ((lids  li(i(l\  whorl  willi  10-12  priuKUA  spir;il 
colds.  ;ib()Mt  4  5  \isiblc  on  penultimate  whorl.  Spinil 
(Olds  sip;n'at('(l  b\  ii;iirow  sjiiices  tluit  iipjiear  ;is  iiKiscd 
lines  iiistc;i{l  of  s])aees  i  figure  24  >.  .-\\i;il  sculpture  ol 
loimded  ribs,  ;il)out  8  in  number  on  the  bo(l\  wlioii  ;m(l 
11  on  the  penultiiiKite  wliori.  SiikiII  nodes  ;ire  present 
where  priin;u"S'  s[)iral  cords  cross  the  axial  ribs.  On  the 
terminal  \:uix  there  is  a  strong  shoukk'r  node.  Shoulder 
rounded  b.ick  to  a  b;ireK  iiii|iresse(l  sutur(>.  Bodx whorl 
loimded  bill  soiiiew  luit  coiistiicted  behind  the  weiik  si- 


p.  Boudiet  ami  H.  Petit.  2002 


Paw  10:3 


phonal  fasciolf.  Onlci-  lip  thin,  weakly  serrati'.  Iniici' pm- 
ti(in  of  outer  lip  with  a  snmoth  romuled  shell'  rnll(iw<'(l 
In  alniut  14  sharp  short  liiations.  ( 'oliiiiiella  Willi  weak 
callns  on  iiuliietural  area.  Ctiluniella  with  three  lolds  that 
do  not  e.xtenil  to  the  outer  edge  of  the  indnetiira.  i'os- 
terior  told  largest.  Two  anterior  folds  situated  011  ends 
of  a  low  shelf-like  platlorin  with  the  anterioiiiiost  fold 
becoming  the  edge  ol  the  short  well-toniied  siphonal 
canal.  Umbilicus  absent.  Shell  color  mottled  white  and 
light  browii.  ("olors  arranged  in  wfak  bands.  Hmwii 
bands  are  located  at  the  shoulder,  just  antf^rior  to  the 
periphery  ami  at  the  base. 

EtyniologA':  The  (ireek  ki/phoina,  meaning  hump, 
with  lelerenci'  to  the  stiiixK  axial  ribs  of  tlie  species.  To 
be  treated  as  a  noun  in  apposition. 

Remarks:  \ij)j>()iiili)lur(i  ctipJutnui  differs  from  \ip- 
jxnifijiliciyi  iiodosivtiricosfi  (Petncli.  1979',  new  combi- 
nation (originalK  in  .Agrz/ri.vi.  in  ha\ing  iimeli  finer  spiiai 
sculpture,  fewer  and  more  wideK  spaceil  axial  ribs,  and 
a  proportionalK-  liroader  shell.  (The  transfer  of  .Agrt?r/.v 
uodosivaricosa  to  Sippoiuijiliird  will  be  discussed  in  a 
future  monograph.)  The  batlnnietrie  distribution  ol  the 
genus  Nipponapluni  is  imelear  due  to  confused  generic 
allocation  of  di-scribed  species,  but  Nippon(ij)lifni  11/- 
plioma  is  apparenth'  the  species  with  the  deepest  occur- 
rence in  the  genus. 

Xippoiiaplicrii  goiiiata  new  species 
(Figiu-es  25-27) 


Type   Material:      IIolot\pe    (IfS.l    iiini    height 
mm  width)  (dd)  (Figure  25)  in  MNHN. 


11,4 


T^•pe  Localit^•:  .South  of  New  Caledonia,  Norfolk 
lliilge.  Banc  A/ieijne  jalso  called  Banc  Antigonia  on  ma- 
rine charts],  2:3°4r  S,  16S°()1'  E,  2S0  m  [SMIB  5,  sta. 
DWTfi], 

Other  Material  Examined:  South  of  New  Caledonia. 
Norfolk  Ridge.  BIOCAL:  sta.  DW65.  24°48'  S,  168°09' 
E,  245-275^ll,  1  dd.— CALSUB:  dive  21,  22°45'  S, 
lfi7°()9'  E,  340  in,  1  K.— CHALCAL  2:  sta.  f^WTf, 
24°42'  S,  ffiS°l()'  E,  230  m,  3  dd.— SMIB  S:  sta. 
D\VT54,  24°46'  S,  1R8°()8'  E.  235-252  m,  1  dd  (Figure 
27).— Sta.  DW157,  24°4fi'  S.  168°08'  E,  251-255  m.  1 
dd.— Sta.  D\M58,  24°4fi'  S,  lfi8°0S'  E,  262-290  m,  1 
dd.— Sta.  DW159,  24°4rV  S,  Ifi8°()8'  E.  241-245  m.  fi 
dd  (Figure  2fi),-^Sta.  D\\T73,  23'41'  S,  168°()0'  E,  234- 
242  m,  1  dd.— Sta.  D\\T74,  23°40'  S,  lfi8°()f'  E,  235- 
240  m,  2  dd.— BERYX  f  f:  sta.  DWMS.  24M8'  S,  lfiS"09' 
E,  250-270  in,  f  dd. 

Description:  Protoconch  smooth,  of  sliglitK'  more 
than  I  whorl,  tliameter  1.00  mm  (F"igure26).  Teleoconch 
of  5  whorls,  spire  high,  spire  angle  of  63-66°.  Stnictnre 
of  strong  lamellate  axial  ribs  reflected  adapertnralK'  and 
much  weaker  uneven  spiral  cords.  Some  axial  ribs  are 
stronger  than  others  ami  form  stronger  \arices.  Axial  ribs 
numlier  about  L3  on  final  whorl  and  14  on  penultimate 


,uid  .iiilepennltiinate  whorls.  I^ibs  regnl.iiK  cou\cx  on 
first  two  whorls  hecoiniiig  distiiictk  angular  at  shoiikl(.M-. 
Spiral  sciil|)tme  ol  about  S  jirimaiA  cords  on  spire  whorl 
ami  about  15  on  bod\  whorl,  each  with  5-8  secondaiy 
cords  in  intenals.  Spiral  cords  cross  the  man\-  rib-like 
growth  lines  loruiing  finek  reticulate  reticulate  micro- 
sculpture  (Figure  271.  Suture  well  impressed.  Bodv 
whoH  slightly  constricted  behind  si])honal  lasciole.  \p- 
ertnre  with  about  17  liiae  within  that  do  not  extend  to 
the  edge  ol  the  outer  lip  hut  extend  deepK  within.  Inner 
lip  lorming  parietal  shield  extending  slightK'  over  the 
narrow  nmbilicns.  Colnmella  onl\-  sliglitK'  concave  with 
three  folds,  tlii'  anterior  two  close  together,  much  like 
one  large  bibncate  fold.  Siphonal  canal  short  but  dis- 
tinct. Shell  white,  mottled  with  lii^ht  brown  bK)tches 
near  the  peiiphciA".  Priniar\  cords  brown,  especiallv 
wliere  the\'  cross  the  axial  ribs  on  the  last  1.5  whorls. 
Some  specimens  are  completeK  white. 

Etymology:  The  Latin  adjective  '^(iiiirilns  i-a.  -iiiii). 
angnl.iled.  in  relereuce  to  the  shell  outline. 

Remarks:  Sippoitdphrni  <^i)uiala  diliers  from  .V.  tcr- 
(iiiuichii  (Habe,  1961  1  In  its  proportionalK  higher  spire, 
narrower  nmbilicns,  and  its  bnmn  maculations.  From  .V. 
parddlis  it  differs  in  having  thicker  ribs  that  are  not  im- 
bricate. It  also  possesses  a  narrow  umbilicus  and  has  a 
smaller  adult  si«'.  The  two  species,  .V.  pardalis  and  .V. 
goniata.  coexist  south  of  Ni'w  Caledonia  but  have  never 
been  taken  together 

LITER./VTURE  CITED 

.•\(laiiis.  II.  and  .\.  \(laiiis.  I  So 4  |in  1S53-5S].  The  genera  of 
Recent  .Molliisca;  arraiii;e(l  uccoiiling  to  their  organiza- 
tion. 2  vols.  John  \aii  \boist.  London.  [1:  f-256,  pis.  1- 
32,  1853;  1:  257-4S4,  2:  1-92,  pis.  33-72,  1854;  2;  9:3- 
284,  pis,  73-96,  1855;  2;  285-112.  pis,  97-112,  1856;  2; 
41.3-540,  pis.  113-128.  1857;  2:  541-660.  pis.  129-138, 
1858.] 

15eii.  .\.  (;.  and  1'  \  Maxwell.  1987.  .\  revision  ol  llie  lossil 
and  living  gastropods  related  to  I'Icsiolrilon  Fischer,  1884 
(Familv  (.'aiicellariidac,  Snhlaniilv  I'lesiotiitoninae  n.  sii- 
bfani.)  with  an  .Appendix:  (ieneni  ol  Bnccinidae  I'isaniinae 
related  to  ('.nluhrnriri  Schiiniacher,  1817.  New  Zealand 
Geologiciil  Siinev  l';ileontoloijit;il  Bnllctin  54:  1-140,  pis. 
1-30. 

Bouchet.  P.  (ed  .  1995.  Hesultals  des  Campagnes  .ML'SOR- 
STOM.  volume  14,  Meiiioires  dii  Mns(''nni  National 
d'llistoire  Naturelle  167:  1-654. 

Bonchet.  P  and  B.  M;nshall  ledsl  2001.  Tropic;il  Oeep-Sea 
Benthos,  volnnie  22,  Menioires  dii  Mnseinii  National 
dllistoire  N;itnrelle  185;  1-106. 

Brocchi.  (.'•.  13.  1814.  Conchiologia  lossile  snhapennina  con  os- 
senazioni  scolopclie  snt;li  .Apennini  e  sni  siiolo  adiacenle. 
Stamperia  Heale.  Milano.  2  vols.  |l;l-240.  2:241-712.  pis. 
1-16.] 

Crosuier  A.  and  V.  Boucliet  (eds).  1991.  Resnitats  des  Cam- 
pagnes MIISORSTOM.  volnmc  7,  Meinoires  du  Mnseum 
National  d'llistoire  Naturelle,  ser  A  150:  1-259. 

Ginelin.  |.  F.  1791.  Ciiroli  a  Linne  Systenia  Naturae  per  regiia 
tria  naturae.  Kditio  deciiiKi  lertia.  \'ol.  1,  pt.  6  (\enncs): 
3021-3910.  Lipsiae. 


Paiie  104 


TIIK  XAl'TILUS.  \ol.  IKS.  No. 


Hahe.  T  1961.  Coloured  illii.stratiims  of  the  sliell.s  oCJapan  (II). 
lioiku.slui,  Osaka.  1S2  pp.,  .Vppeiuli.x  42  pp.,  66  pis.  |.'\ii 
luiglisli  rdiliiin  was  issueil  in  1964.  with  a  lai<;cr  pagi'  si/.c; 
.Shells  ol  the  western  I'aeifie  in  color,  \'ol.  II.  Hoiknsha, 
Osaka.  233  pp..  66  pis.  | 

Haras(>wych,  M.  (;.  and  R.  E.  Petit,  1982.  Notes  on  the  nior- 
pholoi^  ol  Cduccllami  reticulata  (Gastropoda:  C'aneellar- 
iidaet".  The  Nantilus  96:  104-113. 

Ha.segawa.  K.  2()()().  Caneellaiioidea,  pp.  5S1-5S,5.  pis.  2SV)- 
291.  Ill:  T.  Okntani  (ed.).  Marine  Mollusks  in  |apan.  Tokai 
Uni\ersit\  Press,  Tok\Q,  1173  pp. 

Houart,  R.  2001.  Ingensia  gen.  nov.  and  eleven  new  speeies  ol 
.Mnricidae  (Gastropoda)  from  New  Galedonia.  X'annatn. 
and  Wallis  and  Fntnna  Islands.  In:  P.  Bonehet  and  B.  Mar- 
shall (edsl.  Tropical  Deep-Sea  Benthos,  \olnnie  22.  .\le- 
nioires  dn  Mnsenni  National  d'llistoire  Natnrelle.  Paris 
1S5:  243-269. 

Jousseannie,  F.  1887,  La  hmnlle  ili's  Caiicellanulae  i.Moll- 
usques  Gasteropodes).  Le  Natuniliste  9  [  =  ser.  2,  1]:155- 
157,  192-194,  213-214.  221-223. 

.Marshall.  B.  1991.  Mollusca  Gastropoda:  Segnen/.iidae  from 
.New  (Caledonia  and  the  Lovalt\'  Islands.  In:  A.  Grosnier 
and  P.  Bonehet  (edsi.  Resultats  des  Gampagnes  MUSOH- 
.STO.M,  Nolnme  7.  Memoires  dn  Musenm  National 
dTiistoire  Natnrelle,  Paris,  ser.  A  150:  41-109. 

Petit.  R.  E.  1976.  Notes  on  Gancellariidae  (Mollusca:  Gastro- 
poda) -  III.  Tnlane  Stndi(>s  in  Geolog\'  and  Paleontolog\' 
12:  3.3-43.  pis.  1-2. 

Petit,  R.  E.  and  .M.  G.  llarasewvch.  2000.  Three  new  species 
ol  the  genus  Mciica  (Neogastropoda:  Gancellariidae)  troni 
Soutli  Alrica  and  llie  Philippines.  The  Nautihis  1  14:  142- 
148. 


Petuch,  E.  1979.  TweKc  new  Indo-Pacific  gastropods.  Neni- 
ouria23:  1-21. 

Richer  de  Forges,  B.  1990.  Explorations  loi  liath\al  lanna  in 
the  New  (Caledonian  economic  zone.  In:  .\.  (  j'osnier  led.). 
Resultats  des  Gampagnes  MUSORSTOM.  \()lnnie  6.  .Me- 
moires iln  Mnsenm  National  (.rilistoire  Natnrelle.  ser  .\ 
145:  9-54. 

Richer  de  Forges.  B.  1991.  Les  tonds  meuhles  des  lagons  de 
Nou\e!le-Galedonie:  genenilites  et  eehantillonnages  par 
dragages.  Pp.  7-148,  in:  B.  Richer  de  Forges  (ed.).  Le 
benthos  des  fonds  meuliles  des  lagons  de  Non\elle-Gale- 
donie.  volume  1.  Etudes  et  Theses  [ORSTO.M.  Paris]. 

Riclier  de  Forges,  B.  1993.  Champagnes  d'exploration  de  la  fau- 
ne  batlnale  taites  depuis  mai  1989  dans  la  zone  t^cono- 
mique  de  la  Nouvelle-Galedonie.  Listes  des  stations.  In: 
A.  Grosnier  (ed.),  Resultats  des  Gampagnes  MUSOR- 
STOM. \oKmie  10.  Memoires  dn  .Museum  National 
<riIistoire  Naturelle  156:  27-32. 

Richer  de  Forges.  B.  and  G.  Ghe\illon.  1996.  Les  campagnes 
d'echantillonnages  dn  benthos  bath\al  en  Nomelle-Gale- 
donie.  en  1993  et  1994  (B.ATHUS  la  4.  SMIB  8  et  HAL- 
IPRO  1).  In:  A.  Grosnier  (ed.).  Resultats  des  Gampagnes 
MUSORSTOM,  volume  15.  Memoires  dn  Museum  Na- 
tional d'Histoire  Naturelle  168:  33-53. 

Scarabino.  \'.  1995.  Scaphopoda  of  the  tropical  Pacific  and  In- 
dian Oceans,  with  description  of  3  new  genera  and  42  new 
species.  In:  P.  Bouchet  (ed.),  Resultats  des  Gampagnes 
MUSORSTOM,  volume  14.  Memoires  dn  Museum  Na- 
tional d'Histoire  Naturelle  167:  189-379. 

X'erhecken,  .A.  1997.  Mollusca  Gastropoda:  .\ralm'a  Sea  Gan- 
cellariidae collected  durint;  the  K.\Rl'B,\R  cniise.  Me- 
moires du  Museum  National  d'Histoire  N;itnrclle  172: 
295-323. 


THE  NAUTILUS  116(3):105-1()8.  200:2 


Paijc  105 


Spawn  of  the  Patagonian  gastropod  Parcuthnii  pluinhca 
(Philippi,  1844)  (Buccinidae) 


Guiclo  Pastorino 

Miisro  Ar<i;entino  tie  Ciencias 

Xatiirales-CONICET 
A\.  Ancrel  Ckillanlo  470 
C1405  DJR  Buenos  Aires 
ARGENTINA 
r\pastor@cril)a.ed\i.ar 


Pablo  E.  Penchaszadeh 

Miiseo  Anj;entiiu)  de  C^ieneias 
Naturales-C'ONICET 

Aw  Angel  CJallardo  470 

CI405'dJR  Buenos  Aires 

ARGENTINA 

and 

Faeulfad  t\f  Ciencias  Exactas  v 
Naturales 

Universidad  de  Buenos  Aires 

Buenos  Aires 

ARGENTINA 


ABSTRACT 

The  spawn  ot  Piirctithrid  plmnl)cii  (Philippi.  1844)  consists  of 
an  egg  mass  of  4-19  egg  capsules.  The  uncleaved  egg,  in  num- 
ber of  1-5  per  capsule,  measured  about  300  |a.m.  Pdniithrui 
phimbca  has  direct  development  without  niu'se  eggs,  hatclnng 
at  craw  ling  stage.  Extra-vitelline  substances  in  the  intracapsular 
lifjuid  are  likeK'  to  be  the  onK  e.\tra\itelline  tootl  available  for 
the  embrvos.  Hatching  occurs  b\'  nipture  or  dissolution  ol  the 
capsule  wall  in  a  possibly  random  location  (ju  the  (liin  basal 
membrane.  The  number  of  hatchlings  varied  troui  1-5  within 
the  same  spawn.  Hatchling  muuber  and  shell  m/.c  are  related. 


INTHOnUCTION 

The  genus  VdrriilJirid  was  proposcil  hv  Strcliel  (1905) 
in  hi.s  extensive  accoinit  ol  tlie  gastropod  launa  Ironi  the 
Strait  of  Magellan.  He  descrihed  six  new  spe-cies  under 
I'lirciilliriii.  Hovvevvr,  the  validitv  ol  these  species  re- 
iriains  uncertain. 

Pairuthrid  plnmheti  (Philippi.  IS44)  is  the  oldest 
name  of  a  \en'  poKnioiiihic  species  endemic  to  the  .Ma- 
gellanic province  (Figures  1-3).  It  ranges  from  soiitlu'rn 
C.'huhnt  province  {~45°S)  in  Argentina  to  49°S  in  Chile 
(Dell,  1971).  It  is  usually  found  inti-rtidallv  at  lovy  tide 
inider  rocks  and  in  tide  pools.  Ciallardo  and  Penelias/.a- 
deli  (2001)  mentioned  Parciitliha  jihniihiii  as  liaving  di- 
rect or  non-pelagic  development. 

Bnceinids  have  a  wide  range  of  deyelopnicnlal  iiiddes. 
from  tree-.svvimming  planktotrophic  veligt'rs  (Bandel, 
1975,  1976:  Amio,  1963)  to  lack  of  a  lanal  stage  { Fio- 
roni,  1966:  Mattel  et  al.,  19Sfi:  Pdrlniaun,  1925;  Milos- 
lavieh  and  Dnfresne,  1994;  Miloskivicii  aud  Penchasza- 
deh. 1994). 

In  this  note,  we  describe  the  spawn  ol  Ptirculliria 
pluuthi'd  (Philippi,  1S44),  including  the  ninnber  and  size 


of  the  uncleaved  egg.  the  development  pattern  cUid  die 
hatching  stage,  of  a  population  from  Puerto  Deseado, 
Argentina  (—46°  S)  on  the  .\tlantic  coiLst. 

M.\TERIALS  AND  MKTll()i:)S 

Adidts  specimens  and  egg  masses  ol  Parcutliria  phuuhca 
were  collectetl  at  low  tide  in  Dos  Hermanas.  Puerto  De- 
seado.  Santa  ilrnv.  Province,  .Xrgentina  (47°45'S- 
65°55'\^')  during  Fehrnarv  2000.  The  intertidal  area  i.s 
occupied  liv  mvtilids  toiuiing  ininks  on  n)ck"v  platforms. 
Specimens  and  spawns  ol  /'  phiDihca  were  freijuentlv 
found  in  crevices  and  under  rocks.  Adults  specimens  and 
egg  capsules  were  fixed  in  o7(  formalin  on  sea  water. 
Those  with  shelled  embryos  were  presened  in  70%  etli- 
anol.  Most  photographs  of  the  eggs,  capsules  and  em- 
bryos were  taken  using  a  digital  scanning  camera  at- 
tached to  a  Zeiss  Stemi  2000  stereoscopic  microscope. 
All  iiuages  were  digitally  processed. 

RESL ITS 

We  examineil  15  egg  masses,  lor  a  tol.il  number  ol  106 
egg  capsules.  Each  egg  mass  is  composed  ol  4-19  egg 
capsules  (X  =  S.S3,  SD  =  5.07.  n  =  15)  (Figure  6). 
Pareiilhriti  ))liiiiil>cri  has  transparent,  yellowish,  subcir- 
cnlar,  bnlliform  capsules,  which  are  often  aggregated  and 
partialK  ovcilapping  t'ai'h  other  (I'ignres  7.  10),  They  are 
attached  to  the  substrate  bv  a  circular  ba,sal  membrane 
measuring  4.2-5.2  mm  (X  =  4.96,  SD  =  0.22,  n  =  10), 
P'.ach  capsule  has  its  own  basal  membrane  that  is  not 
fused  with  adjacent  ones.  The  ca|)snles  nieasiu'e  3.2-3.S 
nun  in  dianu-ter  (X  =  3.6,  SD  =  O.IS,  n  =  10),  There 
is  a  suture  line  that  splits  the  ca|)snle  in  two  sides  (Fig- 
ures 9,  II).  Tliere  is  no  pre-formed  apical  plate  or  es- 
cape aperture  as  a  plug,  nor  an  escape  slit.  During  iiatch- 


Page  106 


THK  NAUTILUS.  \n 


Ifi.  \o.  3 


Figures  1-9.  I'dirnlhria  phinibi<i  i  rliilippi,  lS44i.  1-3.  MACN  IO.'5i2.  adult  spceiiiicii  IVoiii  Caleta  (.'oylc,  S.iiita  (  ju/,  Ari^riiliiia. 
Scale  l)ar  =  I  (  uj.  4.  Detail  of  tlic  protoconch.  Siali-  liar  =  1  iniii.  5.  Tiirce  ciiibnos  at  tlio  pre-liatciiliiin  staijf.  Scale  liar  =  1 
mm.  6.  Freslily  Ian!  spawn  with  \isilile  cluster  iif  ei^i;s.  Scale  liar  =  .')  iiiiii.  7.  Several  clusters  of  spawns,  the  tup  ca]isnles  with 
three  and  two  einhnns  respcctixcK.  8.  'I'hree  pws  IroMi  llie  s.uiie  capsule.  Seale  liar  ^  oOH  |Jini  9.  Isolaled  capsule  with  five  earl\. 
shelled  enibnos.  Scale  liar  =   1  nun. 


G.  Pastoriiio  and  P.  P].  Pciifliaszadcli.  2002 


Paw  107 


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Figures  10-12.     Egg  capsules  of  rtirciilliiin  plmnhcd  ■  I'liilippi.   1S441.   10.  (.luster  ( if  i •<;<.;  rapsulcs.   II.   Kulatcil  capMilc,   12. 
Middle  section  showing  die  capsule  pnifilc.  Scale  Imis  =   1  nini. 


ing  the  rupture  of  the  liasal  iiu'iiiliianc  tlial  is  imikIi 
thinner  than  the  upper  wall  (li  the  capsule  was  iccmded: 
this  could  he  an  escape  w"a\  for  hatchlings. 

The  uncleaved  eggs  nieasiu'ed  280-340  |xni  in  diam- 
eter (X  =  300,  SD  =  15.6S,  n  =  14)  (P^ignre  S).  They 
are  embedded  in  a  \en  dense  intiacapsiilar  licjnid  that 
becomes,  when  presened  in  ethxl  alcoliol,  opa(jue  and 
flocculates  like  denatured  egg  whites.  This  intra  capsular 
licjuid  is  conipleteK-  translucent  at  the  latest  stages  of 
de\elopnient,  suggesting  that  it  contains  nutritional  sub- 
stances that  would  be  ingested  In  tlie  cmbnos.  \o 
nurse  egjis  were  recorded. 

The  number  of  embnos  was  1-5  within  an\  given 
spawn.  When  fixe  are  present,  one  is  generalK  abortive 
and  is  proliablv  eaten  iiv  the  remaining  embnos.  Hatch- 
ling  number  and  shell  size  are  apparentlv  iclatrd.  When 
a  single  embryo  developed,  the  shell  length  at  tin-  [)ie- 
hatching  stage  was:  2.14—2.19  mm  (X  =  2.1H  nnn,  n  = 
4,  SD  =  ().()3),  with  t\vo  embrvos  the  shell  length  was: 
1.79-2.22  (X  =  1.99  mm,  n  =  13,  SD  =  0.14),  with 
three;  1.60-1.97  (X  =  LSI  mm.  n  =  21,  SD  =  0.09), 
with  four:  0.79-1.S4  mm  (X  =  1.62  nun,  n  =  .33,  SD  = 
0.17'.  We  did  not  find  capsules  at  the  late  pre-hatching 
stage  with  five  embnos.  The  embnos  hatch  as  crawling 
juveniles,  with  a  well-developed  loot  (Figure  .5). 

The  protoconch  has  'IVi-IVa  convex,  somewhat  glo- 
bose, wliorls  (Figure  4).  Whorls  are  completelv  smooth 
and  browmish  in  color  There  is  a  gradual  transition  to 
teleoconch.  Therefore,  changes  in  the  axial  firnamenta- 
tion  are  nsualK'  the  onlv  wav  to  recognize  features  of  the 
adult  shell.  The  first  teleoconch  wiiorl  has  weak  and  ir- 
rcg\ilar  costae  tliat  disappear  graduallv. 

DISCUSSION 

Development  with  planktonic  lanae  has  been  described 
for  several  species  within  the  familv  Buccinidae.  gener- 


alK from  ti'opical  regions  (e.g..  Visdnia  ptisio  (Linnaeus. 
175S)  and  Pitsia  fiiicfa  (Onnad,  1S4B)  (formerlv  in  the 
genus  Caiithani.s):  Bandel,  1975,  1976).  In  temperate 
and  colder  waters  thi're  are  also  examples  of  hatching  of 
veliger  lanae  (e.g.,  Bdhtjlonui  japoiiicti  (Reeve,  1S42); 
Amio,  1963;  Hivest,  19S3>.  However  the  most  common 
mode  of  development  within  the  Buccinidae  involves  the 
lack  of  free-swimming  lanal  stag(>s,  hatching  of  crawling 
juveniles,  and  the  presence  nl  muse  eggs.  The  best- 
kniown  examples  are  Biicciiniin  uiidfitniii  Linnaeus.  175S 
(Fioroni,  1966;  Martcl  ct  al.,  i9S(-;:  Portmann,  1925),  B. 
n/aucuui  Bniguiere,  1792  ( Miloslavich  and  Dufresne, 
1994),  Scarlcsia  (lira  (Reeve.  1S46)  (Rivest.  19S3i,  and 
Cuius  stiiui)s()ui  (M()rch.  1S67'  iWcst.  1979i.  anionic  oth- 
ers. 

Other  spei'ies  of  iinecinidac.  such  ;is  F,li<J,iiiti()pliiis  ilii- 
iciitctiis  (Sav,  !S25i.  Ii;ivc  dirccl  development  vvitliout 
nurse  eggs,  h;ililiin',;  at  tlie  crawling  pe(hveliger  stage 
(MiIosla\ich  and  Piiicliaszadcli.  1994).  In  this  latter  .spe- 
cies, albumin  in  the  intiacapsuhu'  Ii(|uid  is  the  most  im- 
portant extra  food  ;ivaihiblc  foi'  the  embnos  (Miloslav- 
ich, 1999).  (iiveii  the  hick  of  nurse  eggs  aTid  tlu'  large 
difference  between  the  nncleavcd  egg  diameter  and  the 
shell  length  at  hatching,  I'firciilliria  plumbed  ])robablv 
shows  the  same  pattern  of  embnonic  Icedinti  on  sub- 
stances of  the  intracapsular  llnid. 

We  found  several  capsules  with  onlv  one  cnibno  in 
an  earlv  stage  of  devi-lopincnt,  and  the  rest  ot  them  at 
a  later  stage.  The  absence  of  empty  shells  inside  the 
capsule  makes  nnlikelv  the  possibilitv  of  cannibalism  ;it 
a  late  developmental  stage.  The  same  was  ob.sencd  in 
Euiiouiophos  uuiciurlus  (Miloslavich  and  Penchaszadeh, 
1994). 

The  lact  tliat  clusters  loinied  bv  several  c'gg  masses 
of  /'  pluinhcd  are  connnon  could  indicate  a  gregaiious 


Page  108 


THE  NAUTllAiS.  \  ol.  lift,  NO.  3 


spawninif  lH'lia\ior  as  demonstiatccl  lor  sexeial  other 
iieogastropotis  (D'Asaro,  1970)  and  partieiilarlv  for  some 
hiRX'inids  siieli  as  E.  iiiiiciiuiiis  ( Milnslax  icli  and  Pen- 
ciuiszadeh,  1994). 

The  egg  capsules  ol  some  hnceiiiids  present  a  pre- 
formed exit  liole  or  plug.  This  is  the  ease,  among  others, 
u\  liucciniun  tindattiDt.  B.  lu/dropliiiniiin  Hancock,  lS4fi 
(Thorson,  1935)  and  all  the  species  of  the  genus  Ci>lii.s 
(studied  by  Thorson,  1935  and  1946  as  Sipho),  including 
C.jejfretjsiaruis  (Fischer,  1868)  (  =  C.  hoivsei  in  Lebour, 
1937).  In  other  buceinids  the  escape  of  hatchliugs  takes 
place  through  a  widi'  slit  that  is  situated  in  difierent  po- 
sitions according  to  the  species  (e.g.,  Neptiinea  antkjua 
(Linnaeus,  1758),  Beringiiis  tiirtoni  (Bean,  1834)),  or  by 
an  aperture  tolkmang  the  suture  line  of  the  egg  capsule, 
as  in  E.  uuiriiictiis  (Milosla\'ich  and  Penchaszadeh, 
1994).  Despite  the  presence  oi  a  sntiu'e  line  in  the  egg 
capsules  of  F  pl\nnhc(t.  the  opening  iloes  not  occur  along 
tliis  line,  but  b\'  rupture  or  dissolution  of  the  capsule 
wall  in  an  apparentK  random  location  on  the  thin  basal 
membrane. 

The  large  egg  capsules  oi  \\>lutiij)sius  muxciiicHs 
(Gmelin,  1791)  and  Pijntlofusiis  dcfoniiis  (Ree\e,  1847) 
do  not  liave  anv  kind  of  exit  hole  (Thorson.  1940).  This 
is  also  the  ease  in  Parcuthiia  pluinljca,  although  its  egg 
capsules  are  clearK  smaller. 

.ACKXOWLKDCMFA'TS 

Special  thanks  are  tine  to  .\.  Pettoxclio  and  A.  Dolcc- 
maseolo  who  pro\ided  hospitalitx  at  Puerto  Deseado. 
Veronica  hanov  kindly  drew  Figiu'es  10-12.  Two  anon- 
ymous reviewers  improved  considerabK  the  original 
manuscript.  The  fieldwork  was  supported  in  part  b\'  Pro- 
ject PICT  No.  01-04321  from  the  National  Agency  lor 
Scientific  and  Technological  Promotion,  Argentina. 

LITERATURE  CTTED 

Aiiiio,  M.  1963.  A  c(>iiipaniti\c  ciiihnologN'  of  inariiic  gastro- 
pods, with  ceoliitiical  coiisiclcratioiis,  ( 'ontiihiition  Iroiii 
the  Sliiiii()iK)seki  lliii\crsit\  of  Fislierics  12:  229-.'3.5S 

Baiidei.  K.  1975.  iMiihiyoiialgchausc  kaiiliisclicr  .\1cso-  iiiul 
Neogastropoden  (Mollii.sca).  Akademie  dcr  \\  isMiisrlial- 
ten  uiid  (ler  Literature,  .Mainz  1:  1-133. 

Bandel,  K.  1976.  Moipliologic  dor  Gelege  niid  okdlogisclic 
Beobachtuiigeii  an  HiiccinaceoM  ((Jastropoda'  aiis  dcr 
siidliclicn  Karihisciien  Sec.  Boriiicr  '/.oologisilic  iit'ilragc 
27:9f>-1.33, 

D'A.saro.  C.  N.  1970.  Egg  capsules  ol  prosohr.uuli  iiHilliisks 
from  Siiufli  Floritia  and  the  Bahauias  and  notes  on  spawn- 
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440. 

Dell,  R.  K,  1971.  Tlie  uiariue  niollusca  ol  the  Ho\al  Soeiet\ 
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Dominion  .\lusi  lun  7:  155-233. 


Fioroui.  V.  1966  Ziu'  moq^hologie  und  euihnogenese  des 
(larnitniktes  un  der  transitorisclien  organe  bei  Prosobran- 
chiern  (.Mollusca.  CJastropodai.  RiAue  Suisse  de  Zoologie 
73:  621-876. 

Ciallardo,  C.  S.  and  R  F.  Pencha,szadeh.  2()()1.  llatehiug  uioik' 
and  latitude  in  marine  gastropods:  revisiting  Tliorson's 
paradigm  iu  tlie  southern  hemisphere.  Marine  Biologv 
138:  547-552. 

Lebour,  M.  1937.  The  eggs  and  ianae  of  the  British  proso- 
branchs  with  special  reference  to  those  living  in  the  plank- 
ton. Journal  Marine  Biological  .Association  of  the  United 
Kingdom  22:105-166. 

Martel,  A.,  D.  H.  Larrivee,  K.  R.  Klem  and  |,  11.  llinnnehnan. 
1986.  Reproductive  cvcle  and  seasonal  teeihng  activity  of 
the  neogastropod  Bucchniin  iiiuliiluiii-  Marine  Biologv  92: 
211-221. 

.\lilosla\ich.  1'.  1999.  Nutritional  value  ol  die  intracapsular  liq- 
uid of  En<!(>niophos  unicincttis  Say,  1825  (Caenogastro- 
poda:  Bnetinidael.  [oimial  of  Molluscau  Studies  65:  522- 
523. 

Miloslavich,  R  and  R  F.  Renchaszadeh.  1994.  Spawn  and  de- 
velopment of  En<^(mio))lti>s  tiiiiciiHiiis  (Sav.  1825)  (Gastro- 
poda: Prosobranchia)  Irom  the  southern  Caribbean  Sea. 
The  Veliger  37:  425^29. 

Miloslavich,  P.  and  L.  Dufresne.  1994.  Development  and  effect 
of  female  size  on  egg  and  juvenile  production  in  the  neo- 
gastropod Bticcinnm  a/iirwum  from  the  Saguenav  Fjord. 
Canadian  Journal  ol  Fisheries  and  Acjuatic  Sciences  5L 
2867-2872. 

Rhilippi,  R.A.  1844-1851.  Abbikhmgen  und  Beschreibuugen 
neuer  oder  wenig  gekaunter  eonein  lieu  herausgegeben,  2. 
Cassel. 

I'oitmanu,  A.  1925.  Der  eiuthiss  der  .Xiihreier  auf  die  Lani'U- 
Euhvickkmg  von  Buccitiiim  und  Puqnira.  Zeitschrif  fur 
Moiphologie  und  Okologie  der  Tiere  3:  526-541. 

Rivest,  B.R.  1983.  Development  and  the  influence  of  nurse  egg 
allotment  on  hatching  size  in  Scarlcsia  dim  (Reeve,  1846) 
(Prosobranchia:  Buccinidae).  Joimial  ol  Fvperimental  Ma- 
riue Biologv  and  Fcologv  69:  217-241. 

Strebel,  H.  1905.  Beitrage  zm'  Kenntnis  dcr  Mcilluskenl:iun,i 
der  Magalhaen-Rrovinz.  3.  Zoologischeu  jahrbiii-heni.  .\b- 
teilmig  fiir  Svstematik,  Ceographie,  und  Biologic  dcr  Ti- 
ere 22:  575-666. 

Tliorson.  C.  1935.  Studies  ou  the  egg-capsules  and  develop- 
nienl  ( il  :uetic  UKuine  piosobranchs.  .Meddelelser  oui 
(iroufuid:  Konniiissioneu  Inr  \  idiiisk:d)(4ige  I  n- 
dcrsogclscr  1  Crouland  100:  1-71. 

Thorson,  C.  1940.  Notes  on  the  egg-ca|)sules  of  .some  North- 
,\tlantie  jirosobranelis  of  the  genus  Trosclwlia.  Clin/so- 
ili>iiiii\.  \'(i/(//(i/)v(s,  Sipliii  and  Tniphon.  Nidenskalige  Med- 
dck'lscr  Ira  Dansk  uatuihistorisk  Forcuing  i  Kobenhavn 
104:251-265. 

Hiorson,  C,  1916,  Keproduetion  and  lanal  development  ol 
D.cnisli  niaruie  bottom  invertebrates  with  special  reier- 
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Meddelelser  Ira  Foniissionen  lor  Danmarks  Fiskeri-og 
1  l,i\  luidersogelsi-r  Seric:  Plankton  4:  1-523. 

Wcsl.  1),  1,.  1979.  Nutritive  egg  (letcrminatiou  in  ('()///^  S////I/)- 
S()//(  I  l'r(isiibr:ui(  lii.i:  Huccimd.ie  i.  \nici'ieaii  Zoologist  19: 
956. 


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